CRABS OF THE GENUS GEOTHELPHUSA STIMPSON, 1858
(CRUSTACEA: DECAPODA: BRACHYURA: POTAMIDAE)
FROM TAIWAN, WITH DESCRIPTIONS OF
25 NEW SPECIESABSTRACT. - Twenty-eight potamid crabs of the genus Geothelphusa are reported from
Taiwan. Of these, 25 are described as new, most of which have restricted distributions andoccur in relatively low altitudes (less than 1000 m above sea level). Of the new species, only
G. olea has a relatively wide distribution in western Taiwan, while G. eurysoma, G. monticola
and G. takuan are found only in altitudes between 1000 to 2000 m above sea level.

Introduction
Material and methods
Taxonomy
Geothelphusa Stimpson, 1858
Key to the Taiwanese species of Geothelphusa
Geothelphusa caesia, new species
Geothelphusa ancylophallus, new species
Geothelphusa chiui Minei, 1974
Geothelphusa albogilva, new species
Geothelphusa wangi, new species
Geothelphusa eucrinodonta, new species
Geothelphusa wutai, new species
Geothelphusa tsayae, new species
Geothelphusa bicolor, new species
Geothelphusa miyazakii (Miyake & Chiu, 1965)
Geothelphusa candidiensis Bott, 1967
Geothelphusa ferruginea, new species
Geothelphusa tali, new species
Geothelphusa dolichopodes, new species
Geothelphusa taroko, new species
Geothelphusa monticola, new species

782
782
783
783
786
789
789
792
794
796
799
801
801
803
805
807
809
811
813
815
815

J.-Y. Shy, H.-P. Yu - Graduate School of Fishery Science, National Taiwan Ocean University, Keelung,Taiwan, Republic of China. P.K.L. Ng - Department of Zoology, National University of Singapore,
Kent Ridge, Singapore 0511, Republic of Singapore.

Geothelphusa gracilipes, new species
Geothelphusa eurysoma, new species
Geothelphusa yangmingshan, new species
Geothelphusa takuan, new species
Geothelphusa /anyu, new species
Geothelphusa nanhsi, new species
Geothelphusa tawu, new species
Geothelphusa lutao, new species
Geothelphusa cinerea, new species
Geothelphusa nanao, new species
Geothelphusa ilan, new species
Geothelphusa olea, new species
Acknowledgements..:
Literature cited
1\

818
820
823
823
825
827
829
829
832
834
837
837
840
840

The freshwater crabs of Taiwan have been studied by a number of workers, notably de
Man (1914), Parisi (1916), Miyake & Chiu (1965), Bott (1967, 1968a, b, 1969, 1970), Minei(1974), and Hwang & Mizue (1985). Six species in four genera and two families, viz.
Potamidae: Candidiopotamon rathbuni (de Man, 1914), Nanhaipotamonformosanum(Parisi,1916), Geothelphusa miyazakii (Miyake & Chiu, 1965), G. candidiensis Bott, 1967, G. chiui
Minei, 1974, and Parathelphusidae: Somanniathelphusa taiwanensis Bott, 1968, are now
known at present from Taiwan. The genus Candidiopotamon had been placed in the family
Sinopotamidae, and Nanhaipotamon in the Isolapotamidae (Bott, 1970), but both families
are here regarded as junior synonyms of the Potamidae (Ng, 1988; Ng & Dudgeon, 1992).
In recent years, the authors and their colleagues have made extensive collections offreshwater crabs throughout Taiwan. On examination of these, 31 species in four genera are
now known to occur in Taiwan. The majority, 28, belong to the genus Geothelphusa Stimpson,
1858, of which 25 are here described as new.
In the present study, all the species belonging to the genus Geothelphusa in Taiwan arerevised.

Most of the specimens were collected by hand, or with nets, traps and spades in various
rivers, streams and springs throughout Taiwan. The specimens examined are deposited in
the Graduate School of Fishery Science, National Taiwan Ocean University (NTOU), Keelung,Taiwan; Taiwan Museum (TMCD), Taipei, Taiwan; Department of Life Sciences, National
Tsing Hua University (CHCD), Hsinchu, Taiwan, and the Zoological Reference Collection(ZRC), Department of Zoology, National University of Singapore.
The measurements provided are of the carapace width and length respectively. The
abbreviations G1 and G2 are used for the male first and second pleopods (gonopds)
respectively. The terms used in this study essentially follow those used by Ng (1988).
For the G1, the ratios used include the total length divided by the carapace length (TVCL), terminal segment length (TUfSL); terminal segment length divided by width (TSVTSW); and synovial membrane length divided by width (SMUSMW). The ratios of the total
G11ength divided by the distal segment length of the G2 (TUFAL) are provided (Table 1).

Table 1. The G1 and G2 ratios of 28 Geothelphusa species of Taiwan. G1, first gonopod; G2, second
gonopod; TL, total length; SW, subterminal segment width TSL, terminal segment length; TSW,
terminal segment width; SML, synovial membrane length; SMW, synovial membrane width; FAL,
distal segment length.
Character

G1

Species
G. caesia
G. ancylophallus
G. chiui*G. albogilva
G. wangi
G. eucrinodonta
G. wutaiG. tsayae
G. bicolor
G. miyazakii
G. candidiensis
G. ferruginea
G. tali
G. dolichopodes
G. taroko
G. monticola
G. gracilipes
G. eurysoma
G. yangmingshan
G. takuan
G.lanyu
G. nanhsi
G. tawu
G. lutao
G. cinerea
G. nanao
G. ilanG. olea

i\

G2

TUSW

TUTSL

TSUTSW

SMUSMW

TUFAL

3.4

5.5

3.0

4.8
6.3
6.0
5.9
6.4

2.2
3.8

5.3
4.4

0.16
0.21

3.2
3.8
3.0
3.4
3.0
2.9
3.3
3.3
3.9
3.5
3.2
3.1
3.3
3.3
2.9
3.0
3.0
3.1
3.4
2.9
3.0
3.3
3.3
3.3
4.1

6.0
8.8
5.2
2.6
2.7
3.0
4.2
2.9
4.0
3.8
3.3

3.4
2.5
2.3

5.6
6.3

2.8
2.4
2.9
2.2
1.9

5.2
7.8
6.7
6.0
6.6
6.2
6.6
6.1
6.4
4.9
5.5

2.1
2.2
2.3

5.0
4.2
3.8
3.3
3.1
3.6
3.5

2.2
3.0
2.4
2.6
3.1
1.9

6.8
7.4

2.1
2.3

7.1
5.1
5.7
6.4

2.6
3.2
3.6
2.7
3.5
3.2
3.2
5.0

2.7
1.9
3.5
2.3

8.0
5.8
7.5

2.6
2.9

0.21
0.12
0.23
0.20
0.13
0.11
0.15
0.14
0.12
0.12
0.19
0.13
0.15
0.12
0.13
0.16
0.12
0.16
0.17
0.15
0.17
0.14
0.17
0.17
0.20

* after Minei (1974)

TAXONOMYFAMILY POTAMIDAE

ORTMANN, 1896

Geothelphusa

1858

Stimpson,

Geothelphusa Stimpson, 1858: 100; Miers, 1886: 214; Ortmann, 1897: 300; Rathbun, 1898: 27; Bott,
1967: 211; Bott, 1970: 154.
Potamon (Geothelphusa) - Rathbun, 1905: 200; Alcock, 1910: 59; Kemp, 1913: 298; Balss, 1937: 167;
Sakai, 1965: 174.

Type species. - Geothelphusa obtusipes Stimpson, 1858, by subsequent designation by
Rathbun (1898).
Diagnosis. - Carapace oval-shaped, appears rounded, dorsal surfaces almost smooth orslightly rugose. Epigastric and postorbital cristae absent or very low. Anterolateral margin
strongly convex, cristate or appears smooth, rounded; epibranchial tooth small, sometimes
absent; Third maxilliped with distinct flagellum, not extending beyond with of merus. Male
abdomen triangular. G110ng, slender or slightly stout, groove for G2 ventral, subterminal
segment very gradually tapering from relatively broad base, terminal segment usually coneshaped, tapered, straight, curved inwards, upwards or outwards, much shorter than one third
length of subterminal segment, synovial membrane distinct. G2 with distal segment shorter
than half length of basal ~egment.
{'

Distribution.

- Taiwan, Ryukyu Islands and Japan.

Remarks. - The genus Geothelphusa was erected by Stimpson (1858) for two species, G.
dehaani (White, 1847) and G. obtusipes Stimpson, 1858. The type species was not indicated.
Rathbun (1898) fixed the type species as Geothelphusa obtusipes Stimpson, 1858. The genus
was characterised by the carapace being rounded, smooth, the anterolateral margins convex,
and the epigastric and postorbital cristae absent or almost so (Stimpson, 1858, 1907).
Geothelphusa has however, frequently been used merely as a subgenus of Potamon. In any
event, with the then understanding of the genus, many more species have been referred it
to it since Rathbun (1898), including species from Africa. Bott (1967, 1970) resurected
Geothelphusa as a distinct genus, redefining it for species with a slender sword-shaped G1,
the groove for the G2 being ventral, and the terminal segment cone-shaped. He recognised
three species and one subspecies, all from Japan, Taiwan and the nearby islands, viz. G.
obtusipes Stimpson, 1858 (Ryukyu Islands), G. dehaani dehaani (White, 1847) (Japan), G.
dehaani candidiensis Bott, 1967 (Taiwan) and G. levicervix (Rathbun, 1898) (Ryukyu
Islands). Minei (1973) revised the Japanese members of this genus, resurrecting Potamon(Geothelphusa) sakamotoanus Rathbun, 1905, as a distinct species. Bott (1967) originallyregarded this taxon as a subspecies of G. dehaani, but he later (Bott, 1970) synonymised itwith G. obtusipes. Minei (1973) recognised a total of eight species in the genus - G.
obtusipes, G. dehaani, G. sakamotoana, G. levicervix, G. tenuimana (Miyake & Minei, 1965)(Okinawa, Ryukyus), G. miyazakii (Miyake & Chiu, 1965) (Iriomote and Taiwan), G.
candidiensis (as a distinct species) and G. aramotoi Minei, 1973 (Okinawa, Ryukyus). Bott
(1970: 155, in addendum) regarded G. tenuimana as a synonym of G. levicervix but Minei(1973) argued for its separation, a view which the present authors fully agree with. Anotherspecies, G. chiui Minei, 1974, was later described from Taiwan. Minei (1968) presented
some general ecological data on Japanese Geothelphusa. Suzuki & Sato (1994) figured G.
dehaani, G. obtusipes, G. sakamotoana, G. levicervix and an undescribed Geothelphusa from
Kagoshima, southern Japan, providing notes on their ecology. Suzuki & Tsuda (1994)
described the Kagoshima species (Geothelphusa exigua) subsequently. Hwang & Mizue(1985) describes and provides detailed figures of what they identified as G. chiui, G.
candidiensis and G. miyazakii from Taiwan, including the male and female abdomens andtheir third maxilliped features. Yamaguchi & Baba (1993) discussed the types of G. dehaani
in his review of the Japanese crabs studied by De Haan.
Shen (1932) reported a male of Potamon (Geothelphusa) dehaani from Shantung(northern China) but his record has not been substantiated. This record was not listed by Bott
(1967, 1970). Shen's figures however, show a species which has a different carapace andG1 from any known Geothelphusa. The carapace of Shen's specimen resembles some

species of Sinopotamon but the G1 tip is very slender and tapering, unlike known
Sinopotamon or other mainland Chinese genera. Bott (1967) lists specimens of Geothelphusa
dehaani from Yentempo and Tsingtao in China, but in his record of distribution, he writes"Kiiste von China (?)" (Bott, 1967: 212), reflecting his doubt about their identities. A detailed
re-examination will probably show that the mainland Chinese specimens belong to anothergenus and species. The genus Geothelphusa is thus not reliably known outside Japan,
Ryukyus and Taiwan.
The records of G. miyazakii and G. candidiensis from the Ryukyu Islands by Minei (1973)
almost certainly do not belong to these species. We have examined specimens from Japanpreviously referred to G. miyazakii and G. candidiensis and they actually represent
undescribed species. These taxa will be described at a later date when the potamid fauna of
the Ryukyus is revised.
The genus Geothelphusa
species (Table 2).

Stimpson, 1858, as presently defined, contains 35 described

Geothelphusa albogilva, new species (Taiwan)
Geothelphusa ancylophallus, new species (Taiwan)
Geothelphusa aramotoi Minei, 1973 (Ryukyus)
Geothelphusa bicolor, new species (Taiwan)
Geothelphusa caesia, new species (Taiwan)
Geothelphusa candidiensis Bott, 1967 (Taiwan)
Geothelphusa chiui Minei, 1974 (Taiwan)
Geothelphusa cinerea, new species (Taiwan)
Geothelphusa dehaani (White, 1847) = Cancer (Thelphusa) berardi De Haan, 1835 (Japan)
Geothelphusa dolichopodes, new species (Taiwan)
Geothelphusa eucrinodonta, new species (Taiwan)
Geothelphusa eurysoma, new species (Taiwan)
Geothelphusa exigua Suzuki & Tsuda, 1994
Geothelphusa ferruginea, new species (Taiwan)
Geothelphusa gracilipes, new species (Taiwan)
Geothelphusa ilan, new species (Taiwan)
Geothelphusa lanyu, new species (Taiwan)
Geothelphusa levicervix (Rathbun, 1898) (Ryukyus)
Geothelphusa lutao, new species (Taiwan)
Geothelphusa miyazakii (Miyake & Chiu, 1965) (Taiwan)
Geothelphusa monticola, new species (Taiwan)
Geothelphusa nanao, new species (Taiwan)
Geothelphusa nanhsi, new species (Taiwan)
Geothelphusa obtusipes Stimpson, 1858 (type species) (Ryukyus)
Geothelphusa olea, new species (Taiwan)
Geothelphusa sakamotoanus (Rathbun, 1905) (Ryukyus)
Geothelphusa takuan, new species (Taiwan)
Geothelphusa tali, new species (Taiwan)
Geothelphusa tawu, new species (Taiwan)
Geothelphusa tenuimana (Miyake & Minei, 1965) (Ryukyus)
Geothelphusa tsayae, new species (Taiwan)
Geothelphusa taroka, new species (Taiwan)
Geothelphusa wangi, new species (Taiwan)
Geothelphusa wutai, new species (Taiwan)
Geothelphusa yangmingshan, new species (Taiwan)

1. Anterolateral margin faintly cristate, cristae sometimes indistinct, with margin appearingsmooth, rounded2
Anterolateral margin distinctly cristate6
2. Epibranchial tooth low to very low, but visible; branchial region distinctly swollen(Kaohsiung, southern Taiwan)
G. eaesia
Epibranchial tooth absent; branchial region gently convex or appears almost flat .. 3
3. Total length of second ambulatory leg 2.0 times carapace length; G1 subterminal
segment strongly curving outwards (Kaohsiung, southern Taiwan).
.................... \'.
G. aneylophallus
Total length of second ambulatory leg less than 2.0 times carapace length; G 1
subterminal segment gently curving outwards, with the tip sometimes directed upwards...................................................................................................................................4
4. G1 subterminal and terminal segments curving outwards (Hsinchu, western Taiwan)........................................................................................................................
G. ehiili
G 1 subterminal segment gently curving outwards, terminal segment slightly curved
inwards or upwards
5
5. Carapace very high, width to height ratio 1.9; G1 terminal segment slender, length
to width ratio about 3.4 (Pingtung, southern Taiwan)
G. albogilva
Carapace width to high ratio 2.0; G 1 terminal segment length to width about 2.5 (Ilan,northeastern Taiwan)
G. wangi
6. Anterolateral margin lined with small rounded granules
7
Anterolateral margin not lined with small rounded granules, or if present, very low andindistinct
14
7. Epibranchial tooth present
Epibranchial tooth absent or very faint810

8. Dorsal surface of carapace flat, width to height ratio 2.3; epibrancial tooth welldeveloped, distinct; outer proximal margin of G 1 subterminal segment with a distinct
tooth (Taipei, northern Taiwan)
G. euerinodonta
Dorsal surface of carapace gently convex, width to height ratio 2.1; epibranchial tooth
small; outer proximal margin of G 1 entire, without tooth or if present, low and indistinct...................................................................................................................................9
9. Anterolateral region covered with well developed, distinct striae; G1 subterminal
segment curving inwards; G2 total length about 5.4 times length of distal segment(Wutai, southern Taiwan)
G. wutaiAnterolateral region covered with fine striae; G1 subterminal segment straight; G2
total length about 7.7 times length of distal segment (Kaohsiung, Pingtung, southern
Taiwan)G. tsayae
10. Dorsal surface of carapace gently convex, width to height ratio 2.1; G1 terminal segment
slender, length to width ratio 2.9 (Hwalien, eastern Taiwan)
G. bieolorDorsal surface of carapace almost flat, width to height ratio 2.2 or more; G1 terminal
segment relatively stout, length to width ratio less than 2.4
11
11. Ambulatory legs lined with long, thin hairs, in life, with distinct orange and purple
banding; G1 S-shaped (Taipei, Keelung and Ilan, northern Taiwan) ... G. miyazakii
Ambulatory legs lined with thick or thin short hairs, in life legs not distinctly banded;
G1 slightly sinuous or almost straight12
12. Dorsal surface of carapace flat, width to height ratio 2.2; total length of second
ambulatory leg about 1.9 times carapace length; G1 terminal segment short, length
to width ratio 1.9 (Taipei to Nantow, western Taiwan)
G. candidiensis
Dorsal surface of carapace almost flat, width to height ratio more than 2.2, total lengthof second am~ulatory leg more than 2.0 times carapace length; G1 terminal segment
length to width ratio more than 2.1
13
13. Carapace width to height ratio 2.3; tip of medium lobe of epistome rounded; total lengthof second ambulatory 2.1 times carapace length; G1 straight (Pingtung, Southern
Taiwan)G. ferruginea
Carapace flat, width to high ratio 2.4; tip of medium lobe of epistome sharp; total
length of second ambulatory leg 2.0 times carapace length; G 1 gently sinuous (Ilan,northeastern Taiwan)
G. tali
14. Ambulatory legs long, total length of second ambulatory legs more than 2.2 times
carapace length15
Ambulatory legs relatively shorter, total length of second ambulatory less than 2.2
times carapace length16
15. Anterolateralregion covered with with fine striae; G1 subterminal and terminal
segments straight, inner proximal margin dilated (Hwalien, eastern Taiwan).
............
G. dolichopodes
Anterolateral region covered with small, low, rounded granules; G1 slightly S-shaped,
inner proximal margin not dilated, terminal segment gently curving outwards (Hwalien,
eastern Taiwan)
G. taroko
16. Ambulatory legs slender, length of second ambulatory merus more than 4.5 times width.................................................................................................................................17
Ambulatory legs relatively stouter, length of second ambulatory merus less than 4.2
times wi dth
19
17. Carapace width to length ratio 1.3; G 1 subterminal segment slightly S-shaped, terminal
segment straight, high montane species (ca. 2000 m above sea level) (Taichung, middle
Taiwan)G. monticola
Carapace width to length ratio 1.4; G1 subterminal segment straight or curving
outwards; montane species (ca. 1000-1500 m above sea level)
1818. G1 straight, total length about 6.4 times length of short terminal segment (Hwalien,
eastern Taiwan)
G. gracilipes
G1 curving inwards, total length about 4.9 times length of slender terminal segment(Taichung, middle Taiwan)
G. eurysoma

19. Ambulatory legs slender, length of second ambulatory merus more than 4.0 times width.................................................................................................................................
20
Ambulatory legs stout, length of second ambulatory merus less than 4.0 times width.................................................................................................................................
21
20. Dorsal surface of carapace smooth, convex, external orbital angle acutely triangular,
sharp; G1 sinuous (Taipei, northern Taiwan)
G. yangmingshan
Dorsal surface of carapace rough, flat, external orbital angle triangular, stout; G 1 straightor slight curving inwards
22
21. Anterolateral margin rough, lined with with ridges or small, rounded granules
Anterolateral margin somewhat rough, with or without faint ridges

23
25

I'.

22. Epibranchial tooth low, faint, anterolateral region of carapace without ridges (Taoyuan,
northwestern Taiwan)
G. takuan
Epibranchial tooth absent, anterolateral region of carapace with ridges (Lanyu,
southeastern island)
G. lanyu
23. Dorsal surface of carapace convex, width to height ratio 2.0, epibranchial tooth faint;
ambulatory legs lined with short, thick short hairs (Tainan, southeastern Taiwan) .
.
G. nanhsi
Dorsal surface of carapace gently flat, width to high ratio 2.2, epibranchial tooth absent;
ambulatory legs lined with thin short hairs
24
24. Outer proximal margin of G 1 subterminal segment with tooth, terminal segment slender,
length to width ratio 2.7 (Taitung, southeastern Taiwan)
G. tawu
Outer proximal margin of G1 subterminal segment entire, without tooth, terminal
segment stout, length to width ratio 1.9 (Lutao, southeastern island)
G. lutao
25. External orbital angle acutely triangular, sharp; G1 straight, inner proximal margin
slightly dilated, terminal segment slender, length to width ratio 3.5 (Hwalien andTaitung, eastern Taiwan)
G. cinerea
External orbital angle triangular, stout; G 1 curving inwards or outwards, outer proximal
margin dilated, terminal segment length to width ratio less than 3.5
26
26. Dorsal surface of carapace almost flat, width to height ratio 2.2; G 1 subterminal segment
almost straight, terminal segment slightly curving outwards
G. nanao
Dorsal surface of carapace convex, width to height ratio 2.0; G 1 subterminal segment
gently curving outwards, terminal segment gently curving outwards or inwards 27
27. Distance between tip of male abdomen and anterior margin of sternite 4 about 2.0
times that of length of sternites 1-3; G1 terminal segment gently curving outwards,
ratio of total length to terminal segment length 5.8, synovial membrane length towidth ratio 3.2
G. ilanDistance between tip of male abdomen and anterior margin of sternite 4 about 1.4
times that of length of sternites 1-3; G1 terminal segment slight curving inwards,
ratio of total length to terminal segment length 7.5, synovial membrane length towidth ratio 5.0 (Taipei to northern Kaohsiung, western Taiwan)
G. oleaGeothelphusa caesia, new species(Fig. 1)
MateriJll examined. - Holotype - Male, 32.2 by 24.8 rom (NTOU FlO022), KAOHSIUNG HSIEN:
Chiahsien, colI. J.Y. Shy & W.L. Tsay, 6.viii.1992.
Paratypes - KAOHSIUNG HSIEN: Chiahsien - 2 females (NTOU FlO018), colI. H.P. Yu,
13.vii.1982; 1 male, 2 females (NTOU FlO019), colI. H.P. Yu, 17.iv.1983; 1 male, 1 female (NTOU
F10021), colI. J.Y. Shy & W.L. Tsay, 6.viii.1992.
Others - KAOHSIUNG HSIEN: Meinung - 1 male (NTOU F10020), colI. J.Y. Shy & P.H. Ho,
8.v.1992; 1 female (TMCD 2845), colI. J.T. Lin, 9.viii.1992; 3 males (TMCD 2846), colI. J.T. Un,
26.xi.1992; 1 male (TMCD 2849), colI. J.T. Un, 27.v.1993.

Diagnosis. - Carapace smooth, anterolateral margin with faint, smooth crista and verysmall epibranchial tobth. Cervical groove shallow, faint. Branchial region flat. Carapace
length 1.6 times longer than deep. Tip of medium lobe of epistome stout. Sternites 1-3 long,
about 1.9 times length between tip of male abdomen and anterior margin of sternite 4. Fingers
of chela forming wide, oval gape when closed. Second ambulatory leg 2.0 times carapace
length. G1 almost straight to slightly curved outwards; base of subterminal segment relatively
narrow, total length about 3.8 times basal width; terminal segment almost straight, with
spinules on distal one-sixth (Fig. 1c-e).

Size. - Largest male 33.8 by 26.5 mm (TMCD 2846); largest female 37.3 by 28.0 mm(NTOU FlO019); smallest mature female 29.5 by 23.7 mm (NTOU FlO021).

Remarks. - This species is allied to G. albogilva, but it distinguished as follows: 1. thebranchial region of the carapace is flatter and the cardiac region is distinctly flattened; 2.
the G1 subterminal segment is only slightly curved outwards and the synovial membrane
length is about 5.3 times the width; and 3. the G2 distal segment is shorter, the total length
being about 6.2 times the distal segment length (4.8 times in G. albogilva).
Etymology. - The name is derived from the Latin for bluish-gray, with reference to thecarapace colour of the adult crabs.

Geothelphusa ancylophalius, new species(Fig. 2)
MateriJll examined. - Holotype - Male, 32.4 by 25.6 mm (NTOU FlO117), KAOHSIUNG HSIEN:
Neimen, Mucha, colI. J.Y. Shy & P.H. Ho, 8.v.1992.
Paratypes - KAOHSIUNG HSIEN: Neimen, Mucha, 1 male, 1 female (NTOU FlO023), colI. J.Y.
Shy & P.H. Ho, 8.v.1992; 3 males, 1 female (NTOU F10024), colI. J.Y. Shy & P.H. Ho, 9.v.1992.
Others - TAlNAN HSIEN: Nanhsi -1 male (NTOU FlO025), coIl. J.Y. Shy & P.H. Ho, 7.viii.1992.

Diagnosis. - Carapace smooth, strongly convex transversely and longitudinally, carapace
length about 1.5 times depth. Anterolateral margin with smooth crista, without epibranchial
tooth. Cervical groove shallow, faint. Branchial region higher than gastric region. DistanceFig. 1. Geothelphusa caesia, new species. Holotype male, 32.2 by 24.8 mm. a: dorsal view; b: frontal
view; c: ventral view ofright Gl; d: dorsal view of right Gl; e: terminal segment of Gl; f: ventral viewof right G2. Sc~le = 1.0 mm.

Fig. 2. Geothelphusa ancylophallus, new species. Holotype male, 32.4 by 25.6 mID. a: dorsal view;b: frontal view; c: ventral view of right Gl; d: dorsal view of right Gl; e: terminal segment of Gl; f:ventral view of right G2. Scale = 1.0 mID.

between tip of male abdomen and anterior margin of sternite 4 about 1.8 times of sternites
1-3. Fingers of chela forming large gape when closed. Male abdominal segment 7 short,width about 1.5 times length. Gl strongly curved upwards; terminal segment long, straightor slightly curved outwards, total length about 4.8 times length of terminal segment; terminal
segment length about 3.8 times width (Fig. 2c-e).

Size. - Largest male 32.4 by 25.6 mm (NTOU FlO1l7); largest female 29.9 by 24.2mm(NTOU Fl0023); smallest mature female 24.8 by 19.3 mm (NTOU FlO024).
i'.

Distribution. - Southwestern Taiwan.
Remarks. - This species is allied to G. albogilva and G. caesia, but it can distinguishedas follows: 1. the second ambulatory legs is longer, being about two times the carapace length,with the merus being more rounded in cross-section, the width about 1.6 times the height(1.8 times in G. albogilva and G. caesia); 2. the male abdominal segment 7 is
proportionately shorter; and 3. the Gl is strongly curved outwards, the terminal segment is
proportionately longer and the length of the synovial membrane is about 4.4 times that of
the width.
Etymology. - The name is derived from the Greek "ankylos" (for bent) and "phallus" (forgonopod), alluding to the distinctly bent Gl of the species.

Geothelphusa chiui Minei, 1974
(Fig. 3)
Geothelphusa
Geothelphusa

chiui Minei, 1974: 243, figs. 4,5; 6E, F [type locality: Hsinchu Hsien, Taiwan].
chiui - Hwang & Mizue, 1985: 13 (part), text fig. 8, pI. lIB.

Diagnosis. - Carapace smooth, strongly convex transversely and longitudinally.
Anterolateral margin smooth. Fingers of chela forming wide gape when closed. Ambulatory
legs slender, smooth, with very short hairs; second ambulatory leg 1.4 times as long as broad;dactylus slender. Male abdominal segment 6 0.55 times as long as broad; segment 7 0.78
times as long as broad, 1.15 times length of segment 6. Gl moderately curved outwards;
subterminal segment length 6.3 times that of terminal segment; terminal segment slender,
straight; synovial membrane 6.0 times as long as broad (Fig. 3b, c; from Minei, 1974).

Fig. 3. Geothelphusa chiui Minei, 1974. Holotype male, 34.0 by 26.7 mm. a: dorsal view; b: dorsal
view of G1; c: ventral view of Gl. (After Mmei, 1974).

Remarks. - The identity of this species causes some problems as we have not been able
to find specimens in Taiwan exactly referrable to it. Minei (1974) described the holotype
from a town called Nanpu in Hsinchu Hsien (northwestern Taiwan) and provided good
descriptions and illustrations of the species, including its G 1. Fresh collections made in andaround Nanpu have not uncovered this but another species instead, G. olea, new species.
Geothelphusa olea differs from G. chiui .in several aspects, viz. the carapace is more
flattened (against distinctly swollen in G. chiui), the anterolateral margin of the carapace isdistinctly cristate and granulated (indistinctly cristate and smooth in G. chiui), the larger malechela of adult males does not have a large gape when the fingers are closed (with a very wide
gape in G. chiui) and the Gl terminal segment gently curves upwards (curves outwards in
G. chiui). We have not been able to find any species in the Hsinchu area which has the
inflated physiognom?, chafacteristic of G. chiui s. str.
\.

The holotype male and one female were supposedly collected by a parasitologist, Dr.
Chiu Jui-Kuang in December 1960 (Minei, 1974: 243) but no other data is available. It isinteresting to note that in Chiu's (1964) paper detailing his crab collections and theirassociation with Paragonimus, there were no records from Nanpu in Hsinchu Hsien. It is
possible that the specimens were actually collected from other areas and had been incorrectly
labelled.
We have obtained three other new species (G. albogilva, G. ancylophallus and G. wangi)
which also have a similar physiognomy to G. chiui, but their GIs all differ from that of G.
chiui substantially (see Minei, 1974: Fig. 6E, F). Only in G. chiui is the Gl terminal segment
gently curving outwards.
The identities of the other specimens reported by Minei (1974: 243) from Kuanhsi (Hsinchu
Hsien), Hsin-I (Nantow Hsien) and "Taiwan" as "G. chiui" will have to checked to ascertain
their identities. It is possible that these specimens contain more than one species. We havenot been able to obtain the type specimens of G. chiui as they are not easily accessible. The
collections of the Zoological Laboratory of the Kyushu University, where the specimens
were originally deposited, have been transferred to the Kitayushu Museum of Natural History(K. Baba and M. Takeda, pers. comm.), and much of the crab collections have not been
properly arranged and cannot be easily located without a detailed search.
Once the types are available, it would be best to redescribe in detail G. chiui and sort out
the actual identities of Minei's other specimens. For the moment, we feel that it is best torecognize G. chiui as a distinct species.

Geothelphusa

albogilva, new species(Fig. 4)

Geothelphusa chiui - Hwang & Mizue, 1985: 13 (part), text fig. 8, pI. lIB (nee Geothelphusa
Minei, 1974)

chiui

Material examined. - Holotype - Male, 42.1 by 32.9 mm (NTOU FI0189), PINGTUNG HSIEN:
Hengehun, Shehding, colI. lY. Shy & P.H. Ho, 19.xii.1990.
Paratypes - PINGTUNG HSIEN: Hengehun, Shehding - 1 male, 2 females (NTOU FIOOO1), colI.
J.Y. Shy & P.H. Ho, 19.xii.1990; 1 male, 1 female (NTOU FI0003), colI. lY. Shy & P.H. Ho, 22.i.1992;
1 male, 2females (NTOU FI0008), colI. J.Y. Shy & P.H. Ho, 23.i.1992; 1 male, 5 females [1 ovigerous 1(NTOU FIO006), colI. lY. Shy & P.H. Ho, 7.v.1992. - Hengehun, Byitztou - 1 male, 2 females

Fig. 4. Geothelphusa albogilva, new species. Holotype male, 42.1 by 32.9 mm. a: dorsal view; b:frontal view; c: ventral view of right Gl; d: dorsal view of right Gl; e: terminal segment of Gl; f:ventral view of right G2. Scale = 1.0 mm.

(NTOU F10007), J.Y. Shy & W.L. Tsay, 4.viii.1992. - Chialuohshoei -1 male (NTOU FlO014), coli.
J. Y. Shy & P. H. Ho, 22.i.1992. - Kending - 1 male, 2 females (NTOU FlO012), coli. J.F. Hwang,
24.vi.1988. - Manchow, Kangkou - 1 male (NTOU FlO015), coli. J.Y. Shy, 7.vii.1990. - Manchow,
Jeoupemg - 1 female (NTOU FlOOl3), colI. J.Y. Shy & P.H. Ho, 22.i.1992; 2 females (NTOU FlO017),
coli. P.H. Ho, 19.iii.1992. - Manchow, Nanren Lake - 1 male, 2 females (NTOU F10004), coli. J.Y.
Shy, 7.vii.1990. - Suchongshi, - 12 males, 12 females (NTOU FlO002), coli. H.P. Yu & U. Hwang,
14.vii.1982; 1 male (NTOU F10016), colI. J.Y. Shy & H.G. Lai, 11.x.1986; 2 males, 1 female (NTOU
F10005), 1 male, 1 female (ZRC 1994.4221), coli. J.Y. Shy, 7.vii.1990; 1 male (NTOU FlOOll), colI.
J.Y. Shy & P.H. Ho, 6.v.1992. - Wutai - 4 males, 1 female (NTOU FlO010), colI. H.P. Yu & U.
Hwang 12.vii.1982.
TAITUNG HSIEN: Tawu - 1 male, 1 female (NTOU F10009), coli. H.P. Yu & J.J. Hwang,
29.xii.1982.

Diagnosis. - CaraJ\llce smooth, carapace length 1.5 times longer than depth, strongly convex
transversely and longitudinally. Anterolateral margin with faint, smooth crista, without
epibranchial tooth. Cervical groove very shallow, faint. Tip of medium lobe of epistome
stout. Fingers of chela forming very wide gape when closed. Second ambulatory leg short,about 1.7 times carapace length. G1 subterminal segment gently curved outwards; outerproximal margin with a tooth, inner proximal margin dilated; distal part of G 1 terminal segment
slightly curved upwards (Fig. 4c-e).

Size. - Largest male 42.1 by 32.9 mm (NTOU F10189); largest female 48.3 by 36.7 mm(NTOU FlO002); smallest mature female 26.7 by 21.0 mm (NTOU FlO015).

Remarks. - This species is allied to G. wangi, but it distinguished by the following
aspects: 1. the cristate anterolateral margin is smooth, without granules; 2. the male abdominal
segment 6 is proportionately longer, the length to width ratio is 2.3; 3. the G1 terminal segment
is slender and curving inwards, with the length to width ratio 3.4; and 4. the G2 distal segment
is long, with the total length to distal segment length ratio 0.21.
Etymology. - The name is derived from the Latin for yellowish-white, with reference tothe distinctive carapace colour of adult crabs.

Geothelphusa wangi, new species(Fig. 5)Material examined. - Holotype - Male, 29.0 by 23.9 mm (NTOU F10197), ILAN HSIEN: Chilan,
coli. C.H. Wang, 11.xi.1981.
Paratype - ILAN HSIEN: Chilan - 1 female (NTOU FlO186), colI. C.H. Wang, 11.xi.1981.

Diagnosis. - Carapace smooth, convex transversely and longitudinally, carapace length
1.5 times longer than depth. Anterolateral margin with faint, smooth crista, lined with small,very low granules, without epibranchial tooth. External obi tal angle sharp. Distance between
tip of male abdomen and anterior margin of sternite 4 about 1.9 times length of sternites 13. Fingers of chela forming very wide gape when closed. Male abdominal segment 7 bell-

Fig. 5. Geothelphusa wangi, new species. Holotype male, 40.5 by 31.0 mm. a: dorsal view; b: frontal
view; c: ventral view of right Gl; d: dorsal view ofright Gl; e: terminal segment of Gl; f: ventral viewof right G2. Scale = 1.0 mm.

Fig. 6. Geothelphusa eucrinodonta, new species. Holotype male, 18.5 by 13.8 mm. a: dorsal view;b: frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment of G1; f:ventral view of right G2. Scale = 1.0 mm.

shaped and short, width about 1.5 times length. G1 subterminal segment gently curved
outwards, outer proximal margin with a tooth, inner proximal margin dilated; terminal segment
cone-shaped, almost straight (Fig. 5c-e).

Size. - Holotype male 29.0 by 29.3 mm (NTaD FlO197); female 41.3 by 32.7 mm (NTaD
FlO186).

Distribution. - Northeastern Taiwan.
i'Remarks. - This species is allied to G. albogilva. The distinguishing features of thespecies have already been discussed under the Remarks for G. albogiva.
Etymology. - The species is named after Mr. Wang Chia-Hsiang of the Taiwan Provincial
Museum for all his help during this study.

Geothelphusa eucrinodonta, new species(Fig. 6)
Material examined. - Holotype - Male, 18.5 by 13.8 mm (NTaD FlO199), TAIPEI HSIEN: Kungliao,
Homei, colI. J.Y. Shy & W.L. Tsay, 3.ix.1993.
Paratypes - TAIPEI HSIEN: Kungliao, Homei -1 males, 3 females (NTaD F10097), colI. J.Y. Shy& W.L. Tsay, 3.ix.1993; 1 male (NTaD FlO099), lY. Shy, 29.iv.1992; 2 males, 3 females (NTaD
FlO098), colI. C.S. Heh, 28.vii.1993.

Diagnosis. - Carapace flat, with fine concave pits, width and length about 2.3 and 1.7 times
depth respectively. Gastric region smooth, anterolateral region rough, with fine striae.
Anterolateral crista distinct, lined with granules, epibranchial tooth present. Distance between
tip of male abdomen and anterior margin of sternite 4 about equal to length of sternites 13. Length of second ambulatory leg about 1.9 times carapace length; length of dactylus about
1.1 times that of propodus. Male abdominal segment 7 slightly bell-shaped. G1 relatively
short; subterminal segment straight or slightly curved outwards, outer proximal margin witha tooth; terminal segment straight or slightly curved outwards, spinules on about distal onefifth (Fig. 6c-e).

Coloration. - Carapace brownish-green to orangish-green. Chelae orangish-yellow withdark brown spots. Ambulatory legs light brownish-green with dark-coloured spots.

Size. - Largest male 18.1 by 13.7 mm (NTaD FlO199); largest female 19.6 by 15.5 mm(NTaD

F1(098); smallest mature female 19.6 by 15.5 mm (NTaD FlO098).

Remarks. - This species is allied to G. lanyu, but it distinguished by the following
aspects: 1. the carapace is flatter than G. lanyu and the anterolateral margin has a distinct

Fig. 7. Geothelphusa wutai, new species. Holotype male, 26.0 by 20.8 mm. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 rom

epibranchial tooth; 2. the distance between the tip of the male abdomen and anterior marginof sternite 4 is equal in length to sternites 1-3 (shorter in in G. lanyu); and 3. the G1 terminal
segment is curved outwards (curved inwards and only has fine setae on the distal part in G.

lanyu).
Etymology. - The name is derived from the Greek "eukrinos" (for distinct) and "odontus"(for tooth), alluding to the particularly distinct anterolateral tooth present on the species.

Geothelphusa wutai, new species(Fig. 7)
Material examinedi'- Holotype - Male, 26.0 by 20.8 mm (NTOU 10197), PINGTUNG HSIEN:
Wutai, Ila, colI. H.P. Yu.
Paratypes - PINGTUNG HSIEN: Wutai, Ila - 6 males, 10 females (only one mature) (NTOU 10081),
colI. H.P. Yu.

Diagnosis. - Carapace flat, length about 1.7 times depth. Gastric region smooth,anterolateral region rough, with distinct striae. External orbital angle sharp. Anterolateralcrista distinct, lined with small, low granules, epibranchial tooth small but distinct. Cervicalgroove shallow, faint. Fingers of chela forming oval gape when closed. Male abdominal
segment 6 about 0.9 times length of segment 7. G1 subterminal segment slightly curved
inwards, outer and inner proximal margin dilated, but without tooth; synovial membrane'
length about 2.7 times width; distal part of terminal segment especially slender, almost straight,
spinules on about distal one-third (Fig. 7c-e).

Size. - Largest male 26.0 by 20.8 mm (NTOU F10197); largest female 20.3 by 16.3 mm(NTOU F10109).

Remarks. - This species is allied to G. tsayae, but it distinguished by the following
aspects: 1. the dorsal striae on the anterolateral region are stronger and more distinct;'2. theG1 subterminal segment is slightly curved inwards (straight in G. tsayae); and 3. the total
length of the G2 is about 5.1 times that of the distal segment (7.7 times in G. tsayae).
Etymology. - The species is named after the type locality, and the name is used as a noun
in apposition.

Geothelphusa tsayae, new species(Fig. 8)
Geothelphusa miyazakii - Hwang & Mizue, 1985: 13 (part), text fig. 10, pI. IIC (nee Potamon(Geothelphusa) miyazakii Miyake & Chiu, 1965).
Material examined. - Holotype - Male, 26.5 by 20.6 mm (NTOU F10082), KAOHSIUNG HSIEN:
Sanmin, colI. W.L. Tsay & J.Y. Shy, 6.viii.1992.

Fig. 8. Geothelphusa tsayae, new species. Holotype male, 26.5 by 20.6 mm. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 mm.

Paratypes - KAOHSIUNG HSIEN: Meinung - 11 males, 3 females (NTOU FlO085), 1 male, 1female (ZRC 1994.4222), colI. J.Y. Shy & P.H. Ho, 8.v.1992.
Others - KAOHSIUNG HSIEN: Neimen, Mucha - 1 male (NTOU FlO084), colI. J.Y. Shy & P.H.
Ho, 8.v.1992. - Meinung - 2 males (TMCD 2859), colI. J.T. Lin, 26.xi.1992. - Chiahsien - 1 male
(TMCD 2860), colI. 27.v.1993.
PINGTUNG HSIEN: Sandeman - 2 females (NTOU Fl0083), colI. S.P. Yeh, iv.1991.

Diagnosis. - Carapace moderately convex, and length about 2.1 and 1.6 times depth
respectively, with fine concave pits. Gastric region smooth, anterolateral region rough, withstriae. External orbital angle sharp, anterolateral crista distinct, lined with small, low granules,
epibranchial tooth small; frontal region faintly divided into 2 lobes. Distance between tipof male abdomen and anterior margin of sternite 4 long, about 1.7 times length of sternites
1-3. Fingers of chela forming oval gape when closed. G1 subterminal segment straight orslightly curved inwatds, outer and inner proximal margins dilated, outer margin with a verysmall tooth; synovial membrane length about 3.0 times width; distal part of terminal segment
especially slender, slightly curved inwards, spinules on about distal one-quarter (Fig. 8c-e).
Outer proximal margin of G2 basal segment dilated, margin straight to slightly concave (Fig.
8f).
Coloration. - Anterior half of carapace dark orange, posterior half light greenish-brown.
Chelae orangish-yellow.
Chelae and ambulatory legs with orange spots.

Size. - Largest male 29.5 by 23.4 mm (TMCD 2860); largest female 28.5 by 22.0 mm(NTOU Fl0083); smallest mature female 24.6 by 19.8 mm (NTOU FlO083).

Remarks. - This species is allied to G. wutai, and the differences between these two species
have been discussed under the Remarks for G. wutai.
Etymology. - This species is named after Miss Winney Tsay of the National Taiwan Ocean
University, Department of Fishery Science, Crustacean Laboratory, for all her help during
this study.

Geothelphusa bicolor, new species(Fig. 9)
Geothelphusa candidiensis - Minei, 1974: 241 (part), text fig. 2, 6-A, B; Hwang & Mizue, 1985: 13(part) test fig. 9, pI. lID (nee Geothelphusa dehaani candidiensis Bott, 1967).

Material examined. - Holotype - Male, 31.4 by 23.9 mm (NTOU FI0191), TAITUNG HSIEN:
Chiben, colI. J.Y. Shy & W.L. Tsay, 25.viii.1992.
Paratypes - TAITUNG HSIEN: Chiben - 2 males, 3 females (NTOU FlO058), colI. J.Y. Shy &W.L. Tsay, 25.viii.1992. - Chiben· 2 males (NTOU Fl(059), colI. J.Y. Shy & W.L. Tsay, 25.viii.1992.
Peinan, Taiping - 4 males (NTOU FI0065), colI. J.Y. Shy & W.L. Tsay, 26.viii.1992.
Others - TAITUNG HSIEN: Tungho - 1 male, 1 female (NTOU FI0062), colI. lY. Shy & H.G.
Lai, 15.viii.1992; 1 male (NTOU FlO064), colI. lY. Shy & W.L. Tsay, 26.viii.1992. - Peinan, Tananshi- 8 males, 4 females (TMCD 2855), colI. 4.vi.1993.
HWALIEN HSIEN: Yuli - 3 males (CHCD 134), colI. H.C. Liu, 12.v.1993. Juisui - 1 male

Fig. 9. Geothelphusa bicolor, new species. Holotype male, 31.4 by 23.9 mm. a: dorsal view; b: frontal
view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment ofG1; f: ventral viewof right G2. Scale = 1.0 mm.

(NTOU F10063), coil. J.Y. Shy & W.L. Tsay, 27.viii.1992. -Kwungfu- 1 male (NTOU FlO061),
colI. J.Y. Shy & K.Lee, 6.xi.1992. Shoufeng, Takangkou - 1 female (NTOU F10060), colI. J.Y.
Shy & K. Lee, 7.xi.1992.

Diagnosis. - Caroapace smooth. Anterolateral crista distinct, lined with small granules,
epibranchial tooth present. Anterolateral region with faint striae and granules (distinct insmall individuals).
Cervical groove shallow, faint. Merus of second ambulatory legs
moderately flattened laterally, total length about 2.1 times carapace length, merus length
about 3.9 times width; propodus as long as dactylus. Male abdominal segment 7 bell-shaped.
G 1 moderately curved inwards; outer proximal margin of subterminal segment entire or withvery small tooth, inner proximal margin dilated; terminal segment straight or slightly curved
inwards, with spinules on ~bout distal one-sixth (Fig. 9c-e). Total length of G2 about 8.8
times distal segmentiJength (Fig. 9f).
Coloration. - Anterior half of carapace dark brown, posterior half yellowish-brown. Chelae
light orange. Ambulatory legs with fine brown spots.

Size. - Largest male 31.4 by 23.9 mm (NTOU F10191); largest female 32.8 by 24.9 mm(NTOU FlO058); smallest mature female 22.0 by 16.3 mm (NTOU FlO060).

Remarks. - This species is allied to G. taroko, but it distinguished by the following aspects:
1. the color of the carapace is distinctly separated into two parts, the anterior part being
deep brown whilst the posterior part is yellowish-brown; 2. the male abdominal segment 6
is proportionately wider, the width being about 2.3 times that of the length and 1.2 times thewidth of male abdominal segment 7 (about 2.2 times and 1.3 times respectively in G.
taroko); and 3. the G1 is moderately curved inwards, the outer proximal margin of thesubterminal segment has no tooth and the inner proximal margin is dilated.
Etymology. - The name is derived from the Latin, alluding to the distinctive two-colour
tone of the carapace of adult crabs.

Geothelphusa

miyazakii (Miyake & Chiu, 1965)(Fig. 10)

Potamon (Geothelphusa) miyazakii Miyake & Chiu, 1965: 595, pIs. 13, 14 [type locality: Alilao, Taipei
Hsien, Taiwan]
Geothelphusa miyazakii - Minei, 1974: 242, figs. 3, 6C, D.
Geothelphusa miyazakii - Hwang & Mizue, 1985: 13, text fig. 10, pI. IIC.

Material examined. - TAIPEI CITY: Nankang, Denqshanli -1 male (TMCD 2043), colI. W.Y. Lin,5.viii. 1984.
TAIPEI HSIEN: Shuangchi - 1 male, 1 female (NTOU F 10095), colI. H.P. Yu, l.vi.1982.Chisingshan - 1 female (NTOU F10096), colI. 17.vii.1984. Alilao - 1 male (TMCD 2225), colI. T.S.
Wu, xii.1986. - Yehliou - 1 male, 1 female (NTOU F10215), colI. 30.iv.1993. - Juipin - 1 male
(NTOU F10091), colI. J.Y. Shy & H.G. Lai, 8.v.1993.

Fig. 10. Geothelphusa miyazakii (Miyake & Chiu, 1965). Male, 27.0 by 21.4 mm (NTOD FlO091).
a: dorsal view; b: frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal
segment of G1; f: ventral view of right G2. Scale = 1.0 mm.

KEELDNG CITY: 1 male (NTOD FlO094), colI. P.H. Ho, l.vii.1986. FI0116), D.Y. Wang, l.i.1986.

1 male, 1 female (NTOD

ILAN HSIEN: Chiaochi - 2 males (NTOD FI0092), colI. T.Y. Chan, 7.iii.1985. -1
Fl0093), no data.

male (NTOD

Diagnosis. - Carapace with fine concave pits. Anterolateral region with striae. Distance
between tip of male abdomen and anterior margin of sternite 4 about equal to length ofsternites 1-3. Margins of dactylus and propodus of ambulatory legs lined with strong spinesand long setae. Male abdominal segment 7 long, elongate bell-shaped. Gl distinctly sinuous,
outer proximal margin of subterminal segment entire; terminal segment curved inwards, wi th
spinules on about distal one-quarter (Fig. lOc-e).
{'

Coloration - Carapace purple. Ambulatory legs with purple and orange rings in alternately.

Size. - Largest male 31.3 by 24.5 mm (NTaD FlO092); largest female 33.3 by 26.2 mm(NTaD FlO096); smallest mature female 28.3 by 22.4 mm (NTaD FI0116).

Remarks. - Fresh specimens of this species are. very easy to distinguish from other
Geothelphusa species in Taiwan because of the purple carapace, and the purple ambulatory
legs which have orange rings around them. They can also be distinguished by the long setae
and strong spines present on the margins of the ambulatory dactylus and propodus, as wellas the very sinuous G 1.
Specimens from the Ryukyus referred to G. miyazakii by Minei (1973) seem unlikely tobe this species. The Gl of the Japanese specimens (Minei, 1973: Fig. 9G, H) seem to be
more slender and proportionately longer, and the terminal segment also appears to be
proportionately more elongate.

Geothelphusa

candidiensis Bott, 1967
(Fig. 11)

Geothelphusa dehaani candidiensis Bott, 1967: 212, fig. 12. pI. 53 [type locality: Candidius-See, Formosa(Sun-Moon Lake, Nantow Hsien, Taiwan]; Bott, 1970: 157, pI. 40, figs. 62, 63; pI. 53, fig. 64.
Geothelphusa candidiensis - Minei, 1974: 241, figs. 2, 6A, B; Hwang & Mizue, 1985: 13, text-fig. 9,
pI. lID.

Material examined. - TAIPEI CITY: Neishuangchi - 1 female (NTOD Fl0087), colI. lY. Shy &P.H. Ho, 25.i.1992. - Nankang - 1 male (TMCD 2042), colI. W.Y. Lin, 5.viii.1984.
TAIPEI HSIEN: Hsintine - 1 male, 1 female (TMCD 2835), colI. C.H. Wang, 29.viii.1983; 2 males,
3 females (TMCD 2753), colI. C.H. Wang, 5.L 1992. -Pinglin - 4 females (NTOD FI0088), colI. lY.
Shy, 4.ii.I992. - Chinshan - 3 females [1 ovigerous] (NTOD FI0089), colI. J.Y. Shy, 27.vi.1993.
HSINCHD HSIEN: Hengshan - 3 males, 1 female (TMCD 2663), colI. J.P. Fann, 20.ix.I990.
NANTOW HSIEN: Dihli - 1 male (NTOD Fl0090), colI. J.Y. Shy, 28.ix.1986.
Diagnosis. - Carapace with fine concave pits. Gastric region smooth. Anterolateral regionrough, covered with small granules. Anterolateral crista distinct, lined with small granules,

'-::'

/A
','"

r y'

! ,

f

Fig. 11. Geothelphusa candidiensis Bott, 1967. Male, 28.0 by 21.9 mm (NTOD FI0090). a: dorsal
view; b: frontal view; c: ventral view of right Gl; d: dorsal view of right Gl; e, f: terminal segmentof Gl; g: ventral view of right G2. Scale = 1.0 mrn.

epibranchial tooth small. Fingers of chela forming long, wide gape when closed. G 1 almost
straight, outer proximal margin of subterminal segment almost entire or with a very weak
tooth; terminal segment short, straight or slightly curved inwards, with scattered spinules on
about distal one-third (Fig. Hc-f).

Coloration. - Anterior half of carapace brown, posterior part and ambulatory legs greenishbrown to light orange, chelae yellowish-orange, all with darker coloured spots.

Size. - Largest male 28.0 by 21.9 mm (NTOU FlO090); largest female 24.7 by 18.8 mm(NTOU FlO087); smallesrmature female 17.2 by 13.1 mm (NTOU FlO089).
{'

Remarks. - The identity of G. candidiensis has been rather problematic because it is
supposed to have a very wide distribution in Taiwan (Bott, 1970; Minei, 1974) and has also
been reported from the Ryukyu islands (Minei, 1973). The good series of specimens presently
available, as well as topotypic material from Lake Candidus (= Sun Moon Lake) allows us
to define the species more accurately. The G1 of the holotype as figured by Bott (1967: fig.
12; 1970: pI. 40 figs. 62, 63) is very misleading as it seem to show a very slender, gently
tapering and elongated te.rminal segment. Bott apparently, had not positioned the G1
horizontally and it had been figured from the inner marginal view. From the ventral or dorsal
planes, the G1 terminal segment of the present specimens of G. candidiensis s. str. is very
short, but appears elongate from the inner marginal view. This probably led to workers (e.g.
Minei, 1973) to misinterpret Bott's figures and incorrectly identify their specimens.
The present study also shows that what is generally called "G. candidiensis" in Taiwan
actually consists of several species, including a good number of new ones. Geothelphusa
candidiensis s. str. is perhaps most closely allied to G. nanhsi, but it is most easily
distinguished by the anterolateral region of carapace having small granules. Other differences
have been discussed under the Remarks for G. nanhsi.
Specimens from Japan (Ryukyu islands) referred to G. candidiensis by Minei (1973) are
not this species. We have examined specimens from the Ryukyus and there are two species,
both undescribed. It seems possible that once more collections from the Ryukyus become
available, there might perhaps be more than two species. The new Japanese species will be
described elsewhere at a later date.

Geothelphusa fe"uginea, new species(Fig. 12)Material examined. - Holotype - Male, 16.3 by 12.9 mm (NTOU F10202), PINGTUNG HSIEN:
Manchow, Jeouperng, coll. J.Y. Shy & P.H. Ho, 6.v.1992.
Paratypes - PINGTUNG HSIEN: Manchow, Jeouperng - 1 male (NTOU FlOlOS), coll. J.Y. Shy& P.H. Ho, 6.v.1992. - Manchow, Jeouperng - 3 males, 1 female (NTOU F10107), colI. J.Y. Shy &P.H. Ho, 22.i.1992.
Others - PINGTUNG HSIEN: Manchow, Jeouperng - 1 male (NTOU FlO106), colI. P.H. Ho,
19.iii.1992. - Manchow, Nanren Lake -1 male, 1 female [ovigerous] (NTOU FlO108), colI. J.Y. Shy& c.L. Lee, 7.vii.1992.

Fig. 12. Geothelphusa ferruginea, new species. Holotype male, 16.3 by 12.9 mm. a: dorsal view; b:frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment of G1; f:ventral view of right G2. Scale = 1.0 mm.

Diagnosis. - Carapace with fine concave pits, width and length about 2.3 and 1.8 times
depth. Gastric region smooth, anterolateral region rough, with striae. Anterolateral crista
distinct, lined with small granules. Distance between tip of male abdomen and anterior marginof stemite 4 about equal to length of stemites 1-3. Outer surface of chelae rough, lined withsmall granules; fingers of chela forming long, oval gape when closed. Second ambulatory
leg about 2.1 times carapace length. Male abdominal segment 7 moderately bell-shaped. Gl
subterminal segment straight, outer proximal margin with a tooth; synovial membrane length
about 3.8 times width; terminal segment slightly curved inwards, with spinules on distal onetenth (Fig. 12c-e).

Size. - Largest male 19.6 by 15.7 mm (NTOU F10202); largest female 21.0 by 16.7 mm(NTOU FlO107); smallest mature female 20.3 by 15.9 mm (NTOU FI0108).

Remarks. - This species is allied to G. lutao, but it distinguished by the following aspects:
1. the carapace is flatter than that of G. lutao, and the distance of the tip of the male abdomen
and anterior margin of stemite 4 is about equal to that of the length of stemites 1-3; 2. theGl is more slender, the total length being about 3.9 times that of the subterminal segment(about 3.0 times in G. lutao); and 3. the outer proximal margin of the Gl subterminal
segment has a tooth and the terminal segment is distinctly curved outwards.
Etymology. - The name is derived from the Latin for rust-coloured, with reference to thegeneral colour of the carapace of the species.

Geothelphusa tali, new species(Fig. 13)
Material examined. - Holotype - Male, 14.9 by 11.3 mm (NTOU FI0187), ILAN HSIEN: Tali,colI. J.Y. Shy & W.L. Tsay, 4.ix.I993.
Paratypes - ILAN HSIEN: Tali - 8 males, 7 females (NTOU FI0187), coli. J.Y. Shy & W.L. Tsay,
4.ix.1993.
Diagnosis. - Carapace rough, flat, with fine concave pits, width and length about 2.4 and1.8 times depth respectively. Anterolateral region rough, with striae. Anterolateral crista
distinct, lined with granules, without epibranchial tooth. Length of second ambulatory leg
about 2.0 times carapace length; length of dactylus equal to that of pro pod us. Male abdominal
segment 7 bell-shaped. Gl sinuous, outer proximal margin of subterminal segment with atooth, inner proximal margin dilated; distal part of G 1 terminal segment almost straight orslightly curved inwards (Fig. 13c-e).
Coloration. - Carapace olive to greenish-orange, palm and dactylus of chelae orange;
ambulatory legs yellowish-gray, with scattered, darker-coloured stripes.

Fig. 13. Geothelphusa tali, new species. Holotype male, 14.9 by 11.3 mm. a: dorsal view; b: frontal
view; c: ventral view ofright G1; d: dorsal view ofright G1; e: terminal segment of G1; f: ventral viewof right G2. Scale = 1.0 mm.

Size. - Largest male 14.9 by 11.3 mm (NTOU F10187); largest female 20.6 by 15.6 mm(NTOU F10187); smallest mature female 13.3 by 10.2 mm (NTOU F10187).

Remarks. - This species is allied to G. eucrinodonta and G. gracilipes, but it
distinguished by the following aspects: 1. the carapace is flatter than G. eucrinodonta and
G. gracilipes; 2. there is no epibranchial tooth (present in G. eucrinodonta); and 3. theambulatory legs of this species and G. eucrinodonta are wider than G. gracilipes.
Etymology.
is apposition.

- The species is named after the type locality, and the name is used as a noun

Geothelphusa

dolichopodes,(Fig. 14)

new species

Material examined. - Holotype - Male, 30.1 by 23.2 nun (NTOU FI0194), HWALIEN HSIEN:
HsiuIin, Chingshui, colI. J.Y. Shy & K. Lee, 8.xi.1992.
Paratypes - HWALIEN HSIEN: HsiuIin, Chingshui - 4 males, 2 females (NTOU Fl0070), colI.
J.Y. Shy & K. Lee, 8.xi.1992. - Hsiulin, Shiaochingshui - 4 males, 8 females (TMCD 2852), coli.
P.K.L. Ng & C.H. Wang, 3.vi.I993.
Others - HWAUEN HSIEN: Hsiulin, Chungte - 1 male, 1 female (TMCD 2851), colI. P.K.L. Ng& C.H. Wang, 3.vi.1993. - HsiuIin, Shakatang Hsi - 8 males, 5 females CCHCD 111), colI. H.C. Liu.
Diagnosis. - Carapace smooth, frontal margin divide into 2 lobes, (especially in large
specimens). Anterolateral region with fine lateral striae. Anterolateral crista distinct, smooth,
without epibranchial tooth. Ambulatory legs long, thin, total length of second ambulatory
legs about 2.3 times carapace length, length of merus about 4.4 times width, width about 2.0
times height. Male abdominal segment 7 slightly bell-shaped. Gl subterminal segment
straight, outer proximal margin with a tooth, inner proximal margin dilated; terminal segment
almost straight, spinules on distal one-third (Fig. 14c-e).

Size. - Largest male 31.6 by 23.6 mm (TMCD 2852); largest female 36.6 by 26.8 mm(TMCD 2851); smallest mature female 25.8 by 19.4 mm (TMCD 2852).

Remarks. - This species is allied to G. nanao, but it distinguished by the following
aspects: 1. the distance between the tip of the male abdomen and anterior margin of stemite
4 is about 1.2 times the length of stemites 1-3 (about 1.5 times in in G. nanao); 2. the total
length of the second ambulatory leg is about 2.3 times that of the carapace length (about 2.1
times in in G. nanao); and 3. the total length of the G 1 is about 6.2 times that of the terminal
segment (about 8.0 times in G. nanao).
Etymology. - The name is derived from the Greek "dolichos" (for long) and "podes" (forlegs), alluding to the very long legs of this species.

Fig. 14. Geothelphusa dolichopodes, new species. Holotype male, 30.1 by 23.2 mm. a: dorsal view;b: frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment of G1; f:ventral view of right G2. Scale = 1.0 mm.

Geothelphusa

taroko, new species(Fig. 15)

Material examined. - Holotype - Male, 21.3 by 16.2 mm (NTOU F10192), HWALIEN HSIEN:
Taroko, colI. J.Y. Shy & K. Lee, 5.xi.1992.
Paratypes - HWALIEN HSIEN: Taroko - 2 males (NTOU F10066), colI. J.Y. Shy & K. Lee,
5.xi.1992. - Hsiulin, Loshao - 2 males (NTOU F10067), colI. lY. Shy & K. Lee, 6.xi.1992.
Others - HWALIEN HSIEN: Hsiulin, Chankuang Temple - 19 males, 7 females (TMCD 2853),
1 male, 1 female (ZRC *****), P.K.L. Ng & C.H. Wang, 3.vi.1993.
Diagnosis. - Carapace smooth, with fine concave pits; frontal margin gently sinuous,
appears to be divided into f broad lobes. External orbital angle stout. Anterolateral regionslightly rough, with tine concave pits and small, low granules. Anterolateral crista distinct,
lined with small, low1granules, epibranchial tooth small. Total length of second ambulatory
legs about 2.4 times carapace length. G 1 subterminal segment sinuous, outer proximal margin
with a tooth, inner proximal margin almost straight; terminal segment short, distal part slightly
curved inwards (Fig. 15c-e).
Coloration. - Carapace orangish-red. Chelae orangish-yellow.
brown spots.

Ambulatory legs with fineSize. - Largest male 21.9 by 17.1 mm (TMCD 2853); largest female 27.7 by 21.7 mm(TMCD 2853); smallest mature female 17.7 by 13.9 mm (TMCD 2853).

Remarks. - This species is allied to G. gracilipes, but it distinguished by the following
aspects: 1. the anterolateral region of the carapace is slightly rough but is without striae andhas a small epibranchial tooth; 2. the total length of the second ambulatory leg is about 2.4
times that of the carapace length (about 2.1 times and has more setae in G. gracilipes); and3. the Gl subterminal segment is slightly sinuous and the length of the synovial membrane
is about 4.2 times that of the width (straight and about 3.3 times in G. gracilipes).
Etymology. - The species name is derived from the type locality, Taroko National Park,and is used as a noun in apposition.

Geothelphusa

monticola, new species(Fig. 16)

Material examined. - Holotype - Male, 21.9 by 16.5 mm (NTOU F10204), TAlCHUNG HSIEN:
Hoping, Sheauyeakow, colI. J.Y. Shy & W.L. Tsay, 28.vi.1993.
Paratypes - TAICHUNG HSIEN: Hoping, Sheauyeakow - 1 male, 2 females (NTOU F10112), colI.
J.Y. Shy & W.L. Tsay, 28.vi.1993. - Lishan - 1 male, 1 female (NTOU F101130), colI. J.Y. Shy &W.L. Tsay, 28.vi.1993.
Others - TAlCHUNG HSIEN: Hoping, Shenqguang - 1 male (TMCD 2843), colI. S.K. Chan,
5.vii.1977. - Hoping, Songbor - 1 female (NTOU FlO114), colI. lY. Shy & K. Lee, 30.xi. 1992.
Diagnosis. - Carapace with concave pits, frontal margin divided into 2 lobes. Gastricregion smooth, anterolateral region rough, concave pits present slightly larger than those on

Fig. 15. Geothelphusa taroko, new species. Holotype male, 21.3 by 16.2 rnm. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 rnm.

/..
"
~j;
y"""

Fig. 16. Geothelphusa monticola, new species. Holotype male, 21.9 by 16.5 mm. a: dorsal view; b:frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment of G1; f:ventral view of right G2. Scale = 1.0 mm.

gastric region. Anterolateral crista distinct, lined with low, small granules, epibranchial tooth
low. Fingers of chela forming long, oval gape when closed. Merus of ambulatory legs slender,
merus length of second ambulatory leg about 4.6 times width. Width of male abdominal
segment 6 about 2.7 times length, about 1.4 times of length of male abdominal 7 which isbell-shaped. G1 slightly sinuous, outer proximal margin with a tooth; terminal segment
slender, straight (Fig. 16c-e).

Habitat. - Lives in high altitude habitats (about 2000 m above sea level), under boulders
or in burrows near springs.
Size. - Largest mlile 22.5 by 16.9 mm (NTOD F10112); largest female 23.7 by 17.8 mm(NTOD FlO112); smallest mature female 16.6 by 13.0 mm (NTOD Fl01l3).

Remarks. - This species is allied to G. eurysoma and the differences between them havealready been discussed under the Remarks for G. eurysoma.
Etymology. - The name is derived from the Latin "montis" (for mountain) and "colus"(for preference), alluding to the montane habitat of the species.

Geothelphusa

gracilipes, new species(Fig. 17)

Material examined. - Holotype - Male, 21.6 by 16.0 mm (NTOU FlOI93), HWALIEN HSIEN:
Hsiulin, Lushui, coli. J.Y. Shy & K. Lee, 6.xi.1992.
Paratypes - HWALIEN HSIEN: Hsiulin, Lushui - 1 male, 3 females (NTOU FlO068), coli. J.Y.
Shy & K. Lee, 6.xi.1992. - Hsiulin, Holiu - 2 males, 7 females (NTOU FlO069), colI. lY. Shy &K. Lee, 5.xi.1992.
Others - HWALIEN HSIEN: Hsiulin Lushui - 15 males, 18 females [4 ovigerous] (TMCD 2854),
colI. P.K.L. Ng & C.H. Wang, 3.vi.1993. - Hsiulin, Paiyang - 2 males (TMCD 2858), coli. H.C. Liu,
1O.vi.1993.
Diagnosis. - Carapace slightly wider than most known Taiwanese Geothelphusa species,
width and length about 2.3 and 1.4 times of depth respectively, width about 1.7 times length,with fine concave pits; frontal margin indistinctlydivided into 2 lobes. Gastric region smooth;anterolateral region rough, with small striae and granules. Anterolateral crista distinct, linedwith low, small granules, epibranchial tooth completely absent. Distance between tip ofmale abdomen and anterior margin of sternite 4 about 1.2 times length of sternites 1-3.
Ambulatory legs with dense hairs, merus slender, merus length of second ambulatory leg
about 4.7 times of width. Male abdominal segment 7 bell-shaped. Outer proximal margin of
G 1 subterminal segment with a very distinct tooth, inner proximal margin moderately dilated,
subterminal and terminal segments almost straight (Fig. 17c-e).
Coloration. - Carapace greenish-brown.
brown stripes.

Chelae orange, ambulatory legs with scatteredFig. 17. Geothelphusa gracilipes, new species. Holotype male, 21.6 by 16.0 mm. a: dorsal view; b:frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment of G1; f:ventral view of right G2. Scale = 1.0 mm.

Size. - Largest male 23.3 by 17.5 mm (TMCD 2854); largest female 23.7 by 17.8 mm(TMCD 2854); smallest mature female 15.7 by 12.0 mm (NTOU Fl0068).

Remarks. - This species is allied to G. taroko. The distinguishing features have been
discussed under the Remarks G. taroko.
Etymology. - The name is derived from the Latin "gracilis" (for slender) and "pes" (forlegs), alluding to the slender ambulatory merus of the species.

i\

Geothelphusa

eurysoma, new species(Fig. 18)

Material examined. - Holotype - Male, 21.8 by 15.8 mm (NTOU FlOllO), TAICHUNG HSIEN:
Hoping, Chingshan, coli. J.Y. Shy & K. Lee, 19.x.1992.
Paratypes - TAICHUNG HSIEN: Hoping, Chingshan - 2 females (NTOU FlO111), coli. J.Y. Shy& K. Lee, 19.x.1992.
Diagnosis. - Carapace with fine concave pits, slightly wider than most Taiwanese
Geothelphusa species (similar to G. gracilipes), width about 2.3 and 1.4 times of length anddepth respectively; frontal region indistinctly divided into 2 lobes. Anterolateral crista very
distinct, smooth, epibranchial tooth small. Cervical groove shallow, faint. Distance between
tip of male abdomen and anterior margin of stemite 4 long, about 1.8 times length of stemites
1-3. Merus of second ambulatory leg slender, total length leg about 4.5 times width, widthabout 1.6 times height. Male abdominal segment 6 short, width about 2.5 times length; maleabdominal segment 7 bell-shaped. Subterminal segment of Gl curved outwards; terminal
segment distinctly curved inwards, distal one-third with spinules (Fig. 18c-e).
Coloration. - Anterior two-thirds of carapace reddish-brown, posterior one-third andambulatory legs light greenish-brown, spotted. Upper part of chelae reddish-brown, lower
part orangish-yellow.
Habitat. - Occurs in high altitude habitats (more than 1000 m above sea level), in burrows
near springs.

Size. - Largest male 21.8 by 15.8 mm (NTOU FlOllO); largest female 19.0 by 14.3 mm(NTOU FlOlll); smallest mature female 17.9 by 13.8 mm (NTOU FlOlll).

Remarks. - This species is allied to G. monticola, but it distinguished by the following
aspects: 1. the carapace is wider and deeper in G. eurysoma (the carapace width is about 2.1
and 1.3 times that of the depth and length respectively in G. monticola); 2. the lower regionof the Gl is wider and the lower margin of the Gl has a very low tooth; and 3. the total lengthof the G2 is about 8.5 times that of the distal segment (about 6.6 times in G. monticola).
Etymology. - The name is derived from the Greek "eurys" (for broad) and "soma" (forbody), alluding to the particularly broad carapace of the species.

Fig. 18. Geothelphusa eurysoma, new species. Holotype male, 21.8 by 15.8 mm. a: dorsal view; b:frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment of G1; f:ventral view of right G2. Scale = 1.0 rom.

Fig. 19. Geothelphusa yangmingshan, new species. Holotype male, 31.4 by 24.3 mm. a: dorsal view;b: frontal view; c: ventral view of right G1; d: dorsal view of right G1; e: terminal segment of G1; f:ventral view of right G2. Scale = 1.0 mrn.

Geothelphusa yangmingshan, new species(Fig. 19)
Material examined. - Holotype - Male, 31.4 by 24.3 mm (NTOU FI0185), TAIPEI CITY:
Yangmingshan, coll. H.Y. Cheng, iv.1989.

Diagnosis. - Carapace smooth, anterolateral crest distinct, with small, low granules;
epibranchial tooth very low, indistinct. External orbital angle sharp. Cervical groove shallow,
faint. Distance between tip of male abdomen and anterior margin of sternite 4 about 1.6
times to that of length of sternites 1-3. Fingers of chela forming long, oval gape when closed.
Length of second ambulatory leg about 2.1 times carapace length. G1 sinuous; outer proximal
margin of subterminal segment with a weak tooth, inner proximal margin dilated; terminal
segment gently slender, length about 3.1 times width, distal part gently sinuous (Fig. 19ce).

Remarks. - This species is allied to G. bicolor, but it distinguished by the following aspects:
1. the anterolateral region of the carapace is smooth; 2. the G1 subterminal segment curves
outwards; and 3. the G2 outer proximal margin of basal segment is divided into two lobes.
Etymology. - The species name is named after its type locality, Yangmingshan National
Park, and is used as a noun in apposition.

Geothelphusa takuan, new species(Fig. 20)
Material examined. - Holotype - Male, 16.6 by 12.9 mm (NTOU FI0205), TAOYUAN HSIEN:
Fuhsing, Takuan, coll. J.Y. Shy & W.L. Tsay, 1.xi.1992.
Paratypes - TAOYUAN HSIEN: Fuhsing, Takuan - 1 male, 4 females (NTOU FlO115), coll. lY.
Shy & W.L. Tsay, l.xi.1992.

Diagnosis. - Carapace with concave pits, frontal margin vaguely divided into 2 lobes.
Gastric region smooth, anterolateral region rough, concave pits here slightly larger than thoseon gastric region. Anterolateral crista distinct, lined with small, low granules, epibranchial
tooth small. Distance between tip of male abdomen and anterior margin of sternite 4 about
equal to that of length of sternites 1-3. Fingers of chela forming long, oval gape when closed.
Ambulatory legs slender, total length of second ambulatory leg about 2.2 times carapace
length. Width of male abdominal segment 6 about 2.6 times length. Subterminal segment of
G1 moderately curved inwards, outer proximal margin with a tooth; terminal segment short,
almost straight; total length about 6.8 times of terminal segment length, terminal segment
length about 1.9 times of width (Fig. 20c-e).

Habitat. - Occurs in high altitude habitats (above 1500 m asl), living under boulders andstones of streams.

Size. - Largest male 16.6 by 12.9 mm (NTOU FI0205); largest female 18.9 by 14.2 mm(NTOU FI0115); smallest mature female 14.6 by 11.2 mm (NTOU FI0115).

Fig. 20. Geothelphusa takuan, new species. Holotype male, 16.6 by 12.9 mm. a: dorsal view; b: frontal
view; c: ventral view ofright G1; d: dorsal view ofright G1; e: terminal segment of G1; f: ventral viewof right G2. Scale = 1.0 mm.

Remarks. - This species is allied to G. eurysoma and G. monticola, but it distinguishedby the following aspects: 1. the carapace has a epibranchial tooth (epibranchial tooth is absent
or very small in G. eurysoma and G. monticola respectively); 2. the distance between the
tip of the male abdomen and anterior margin of stemite 4 is about equal to that of the lengthof stemites 1-3 (distinctly longer in G. eurysoma and G. monticola); 3. the Gl is slightly
curved inwards in G. takuan (sinuous and curved outwards in G. monticola and G. eurysoma
respectively); and 4. the outer proximal margin of the Gl subterminal segment has a toothin G. takuan and G. monticola (absent in G. eurysoma).

Etymology. - The species is named after the type locality, and the name is used as a noun
in apposition.

Geothelphusa

lanyu, new species(Fig. 21)

Geothelphusa candidiensis - Minei, 1974: 241 (part), test
candidiensis Bott, 1967)

fig. 2-5, 6 (nee Geothelphusa

dehaaniMaterial examined. - Holotype - Male, 20.2 by 15.8 mm (NTOU FlOI00), TAITUNG HSIEN:
Lanyu, coIl. lY. Shy & K. Lee, 18.iv.1993.
Paratypes - TAITUNG HSIEN: Lanyu - 4 males, 2 females (NTOU FlOlOl), 1 male, 1 female(ZRC 1994.4224), colI. J.Y. Shy & K. Lee, 17.iv.1993.
Others - TAITUNG HSIEN: Lanyu - 2 males (NTOU FlOlO2), colI. RF. Shieh, 22.xii.1992; 1
male, 8 females (TMCD 2848), colI. P.K.L. Ng & C.H. Wang, 23.iv.1993; 9 males, 9 females [1ovigerous] (TMCD 2856), colI. P.K.L. Ng & C.H. Wang, 5.vi.1993; 1 male (NTOU FlOlO3), colI.
RF. Shieh & M.S. Hung, 24.vi.1993.
Diagnosis. - Carapace with fine concave pits. Gastric region smooth, anterolateral regionrough, with fine dorsal ridges. Anterolateral crista distinct, lined with small, low granules,
epibranchial tooth absent. Distance between tip of male abdomen and anterior margin ofstemite 4 short, about 0.6 times of stemites 1-3. Merus of ambulatory legs slender, merus
length of second ambulatory leg about 4.2 times width. Male abdominal segment 7 moderately
bell-shaped. Subterminal segment of Gl straight or slightly curved inwards, outer proximal
margin with a tooth, inner proximal margin dilated; length of synovial membrane length
about 2.6 times width; terminal segment short, straight, lined with only fine hairs (Fig. 21c-e).

Size. - Largest male 20.2 by 15.8 mm (NTOU FlOlOO); largest female 20.7 by 15.8 mm(fMCD 2856); smallest mature female 14.4 by 11.0 mm (fMCD 2848).

Remarks. - This species is allied to G. lutao, but it distinguished by the following aspects:
1. the distance between the tip of the male abdomen and anterior margin of stemite 4 is lessthan that of the length of stemites 1-3 «(longer in G. lutao); 2. the dactylus of the ambulatoryleg is longer than the propodus (about equal in length in G. lutao); and 3. the outer proximal

Fig. 21. Geothelphusa lanyu, new species. Holotype male, 20.2 by 15.8 mm. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 mm.

margin of the G 1 subterminal segment has a tooth and the terminal segment is straight (curved
inwards in G. lutao), with the total length of the G 1 being about 7.4 times that of the terminal
segment (about 5.7 times in in G. lutao).
Etymology. - The species if named after the type locality, and the name is used as a noun
in apposition.

Geothelphusa

nanhsi, new species(Fig. 22)

Material examined. - Holotype - Male, 23.3 by 18.3 mm (NTOU FlOI98), TAINAN HSIEN: Nanhsi,
colI. J.Y. Shy & W.L.i'Tsay, 7.viii.1992.
Paratypes - TAINAN HSIEN: Nanhsi - 2 males, 3 females (only one mature) (NTOU FlO086), colI.
J.Y. Shy & W.L. Tsay, 7.viii.1992.

Diagnosis. - Carapace smooth, frontal margin vaguely divided into 2 lobes. Anterolateralregion rough. Anterolateral crest distinct, lined with indistinct low granules, epibranchial
tooth very small. Distance between tip of male abdomen and anterior margin of stemite 4
about 1.3 times length of stemites 1-3. Length of second ambulatory leg about 2.0 times
carapace length. Male abdominal segment 7 bell-shaped, length of segment 6 about 0.9
times length. Outer proximal margin of G 1 subterminal segment dilated, with very a low
tooth, inner proximal margin straight; terminal segment gently curved inwards, spinules on
distal three-fifths (Fig. 22c-e).

Size. - Largest male 23.3 by 18.3 mm (NTOU FlOI98); largest female 22.6 by 17.4 mm(NTOU FlO086).

Remarks. - This species is allied to G. candidiensis s. str., but it distinguished by thefollowing aspects: 1. the carapace is more convex transversely and longitudinally, with thecarapace width and length being about 2.0 and 1.6 times of the carapace depth respectively;
2. the male abdominal segment 6 is longer, the width being about 2.0 times that of the length(about 2.5 times in G. candidiensis); and 3. the Gl terminal segment is more slender, thelength being about 2.3 times that of the width (about 1.9 times in G. candidiensis).
Etymology. - The species is named after the type locality, and the name is used as a noun
in apposition.

Fig. 22. Geothelphusa nanhsi, new species. Holotype male, 23.3 by 18.3 rom. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 rom.

Geothelphusa tawu, new species(Fig. 23)
Material examined. - Holotype - Male, 18.4 by 14.2 mm (NTOU FI0203), TAITUNG HSIEN:
Tawu, colI. lY. Shy & W.L. Tsay, 24.viii.1992.
Paratypes - TAITUNG HSIEN: Tawu - 3 females (only one mature) (NTOU FI0109), colI. J.Y.
Shy & W.L. Tsay, 24.viii.1992.
Diagnosis. - Carapace with fine concave pits. Anterolateral crista distinct, lined with low,small granules. Gastric region smooth; anterolateral region rough, with fine striae. Distance
between tip of male abdomen and anterior margin of sternite 4 about 1.1 times length ofsternites 1-3. Fingers of .chela forming long, oval gape when closed. Length of second
ambulatory leg about 2.0 times carapace length, dactylus about 1.2 times longer than propodus.
Width of male abdominal segment 7 about 1.2 times length, width of male abdominal segment
6 about 1.4 times width of segment 7. G1 slightly curved outwards, with small tooth on outerproximal margin of subterminal segment; terminal segment slightly curved inwards (Fig.
23c-e).

Size. - Largest male 18.4 by 14.2 mm (NTOD F10203); largest female 16.3 by 12.2 mm(NTOD FlO109).

Remarks. - This species is allied to G. ferruginea and G. lutao, but it distinguished by
the following aspects: 1. the carapace width and length is about 2.2 and 1.7 times that ofthe depth in G. lutao and G. tawu (about 2.3 and 1.8 times respectively in G. ferruginea);
2. the distance between the tip of the male abdomen and anterior margin of sternite 4 about
equal to the length ofsternites 1-3 in G. ferruginea and G. tawu (about 1.6 times in G. lutao);
3. the proportions of the length of the G1 synovial membrane to the width are different (2.7,
3.8 and 3.6 times in G. lutao, G. ferruginea and G. tawu respectively); and 4. the G1 terminal
segment is curved inwards in G. lutao and G. tawu (curved outwards in G. ferruginea).
Etymology.
in apposition.

- The species is named after the type locality, and the name is used as a noun

Geothelphusa

lutoo, new species(Fig. 24)

Material examined. - Holotype - Male, 21.4 by 16.2 mm (NTOU FI0200), TAITUNG HSIEN:
Lutao, colI. P.K.L. Ng & C.H. Wang, 6.vi.1993.
Paratypes - TAITUNG HSIEN: Lutao - 11 males, 3 females (NTOU FlO104), colI. P.K.L. Ng &C.H. Wang, 6.vi.I993. - Lutao - 6 males, 13 females (TMCD 2857),1 male, 1 female (ZRC 1994.4225),
colI. P.K.L. Ng & C.H. Wang, 6.vi.1993.
Diagnosis. - Carapace with fine concave pits. Cervical groove shallow, faint. Gastric regionsmooth, anterolateral region rough, with fine striae. Anterolateral crista distinct, lined with

Fig. 23. Geothelphusa tawu, new species. Holotype male, 18.4 by 14.2 mrn. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 mrn.

Fig. 24. Geothelphusa lutao, new species. Holotype male, 21.4 by 16.2 mm. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 mm.

low, small granules, without epibranchial tooth. Distance between tip of male abdomen andanterior margin of sternite 4 about 1.6 times length of sternites 1-3. Propodus length ofambulatory legs equal to dactylus length. Width of male abdominal segment 6 about 2.4
times length. Outer proximal margin of Gl subterminal segment without or with faint tooth,Gl straight or slightly curved outwards; terminal segment gently curved inwards, about distal
one-seventh with spinules (Fig. 24c-e).

Size. - Largest male 21.4-by 16.2 mm (NTOU FI0200); largest female 23.8 by 17.7 mm(TMCD 2857); smalliest mature female 15.7 by 12.0 mm (NTOU FI0104).

Remarks. - The species is allied to G. lanyu and the differences for these two species havebeen discussed under the Remarks for G. lanyu.
Etymology. - The species is named after the type locality, and the name is used as a noun
in apposition.

Geothelphusa cinerea, new species(Fig. 25)
Material examined. - Holotype - Male, 33.7 by 26.3 mm (NTOU Fl0052), HWAUEN HSIEN:
Juisui, colI. J.Y. Shy & W.L. Tsay, 27.viii.1992.
Paratypes - HWALIEN HSIEN: Juisui - 1 male (NTOU Fl(052), colI. J.Y. Shy & W.L. Tsay,
27.viii.1992. - Juisui - 1 male (NTOU FlO055), colI. J.Y. Shy & W.L. Tsay, 27.viii.1992.
Others - HWALIEN HSIEN: Shoufeng, Shyrtiping - 1 male (NTOU FlO121O, colI. J.Y. Shy & K.
Lee, 26.iv.1986. - Shoufeng, Takangkou - 1 male (NTOU FlO054), colI. J.Y. Shy & W.L. Tsay,
28.viii.1992. - Yuli - 1 male (CHCD 134), colI. H.C. Lin, 12.v.1993.
TAITUNG HSIEN: Tungho - 1 female (NTOU FlO057), colI. lY. Shy & H.G. Lai, 15.viii.1992;
1 male (CHCD 123), colI. H.C. Liu, 1O.v.1993. - Tungho, Dergualao - 2 males (NTOU FlO053), colI.
J.Y. Shy & W.L. Tsay, 27.viii.1992. - Tungho, Taiyung - 1 male, 2 females (CHCD 131), colI. H.C.
Liu, l1.v.1993. - Lichi - 1 male, 4 females [1 ovigerous] (NTOU FlO056), colI. J.Y. Shy & W.L.
Tsay, 27.viii.1992. -Peinan,
Chiafong - 3 males, 1 female [ovigerous] (TMCD 2850), colI. H.C. Liu,
3.vi.1993.

Diagnosis. - Carapace smooth, with fine concave pits. Branchial and gastric regions higher
than other regions, anterolateral region rough, with fine striae; anterolateral crista distinct,
lined with small, low granules, with low, indistinct epibranchial tooth. Distance between tipof male abdomen and anterior margin of sternum 4 about 1.5 times to that of length of sternum
1-3. Gl almost straight, outer proximal margin of subterminal segment with a distinct tooth;terminal segment straight, with spinules at about distal one fifth, total length about 6.4 timesof terminal segment length; synovial membrane length about 3.5 times width (Fig. 25c-e).
Total length of G2 about 6.9 times distal segment length (Fig. 25t).
Coloration. - Carapace bluish-gray, chelae and ambulatory legs grayish, with faint scatteredstripes.

Fig. 25. Geothelphusa cinerea, new species. Holotype male, 33.7 by 26.3 mm. a: dorsal view; b: frontal
view; c: ventral view of right Gl; d: dorsal view ofright Gl; e: terminal segment of Gl; f: ventral viewof right G2. Scale = 1.0 mm.

Size. - Largest male 33.7 by 26.3 mm (NTOU F10052); largest female 32.8 by 25.6 mm(NTOU FlO056); smallest mature female 27.1 by 20.9 mm (NTOU FlO056).

Remarks. - This species is allied to G. caesia, but it distinguished by the following
aspects: 1. the anterolateral region of the carapace is swollen and lined with fine striae; 2.
the anterolateral crista is distinct and lined with small granules; and 3. the G1 is straight, thelength of the synovial membrane being about 3.5 times that of the width.
Etymology. - Thdrtame is derived from the Latin for ashy-coloured, with reference to thecolour of the carapace.

Geothelphusa nanao, new species(Fig. 26)
Material examined .• Holotype - Male, 26.2 by 20.2 mID (NTOU FlO195), lLAN HSIEN: Nanao,
Chinyang, coli. J.Y. Shy & M.S. Hung, 30.iii.1992.
Paratypes - lLAN HSIEN: Nanao, Chinyang - 1 male, 1 female (NTOU FlO071), coli. lY. Shy& M.S. Hung, 30.iii.1992. - Nanao, Chinyang - 3 males, 1 female (NTOU FlO073), coli. lY. Shy& W.L. Tsay, 3.ix.1993; 1 female (NTOU FlO075), colI. J.Y. Shy & M.S. Hung, 15.viii.1991; 1
female (NTOU Fl0074), coli. J.Y. Shy & M.S. Hung, ix.1992.
Others - lLAN HSIEN: Nanao, Chinyang - 1 female (NTOU FlO072), coli. J.Y. Shy, 17.v.1991.
- Nanao, Suhwa Road - 5 males, 8 females [1 ovigerous] (NTOU FlO076), call. Wang. - Nanao- 2 males, 2 females (TMCD 2720), call. G.S. Uu, iv.l991.

Diagnosis. - Carapace with fine concave pits. Cervical groove shallow, faint. Gastric regionsmooth, anterolateral region rough, with fine striae. Anterolateral crista distinct, smooth,
without or with very small epibranchial tooth. Distance between tip of male abdomen andanterior margin of sternite 4 about 1.5 times length of sternites 1-3. Merus of ambulatory
legs slender, merus width of second ambulatory leg about 2.1 times of height. G1 subterminal
segment almost straight, outer proximal margin with a tooth, inner proximal margin dilated;
terminal segment short, curved outwards, total length about 8.0 times of terminal segment
length; length of synovial membrane about 3.2 times width (Fig. 26c-e).
Coloration. - Carapace greenish-brown to orange. Smaller specimens with deep orange
carapace, ambulatory legs covered with dark brown spots.

Size. - Largest male 26.2 by 20.2 mm (NTOU F10195); largest female 30.5 by 22.8 mm(NTOU FlO076); smallest mature female 22.1 by 17.4 mm (NTOU FlO076).

Remarks. - This species is allied to G. dolichopodes, but it can be distinguished by theshorter G 1 terminal segment. Other differences have been discussed under the Remarks for
G. dolichopodes.

Fig. 26. Geothelphusa nanao, new species. Holotype male, 26.2 by 20.2 rom. a: dorsal view; b: frontal
view; c: ventral view ofright Gl; d: dorsal view of right Gl; e: terminal segment of Gl; f: ventral viewof right G2. Scale = 1.0 rom.

Fig. 27. Geothelphusa ilan, new species. Holotype male, 27.8 by 22.2 mm. a: dorsal view; b: frontal
view; c: ventral view ofright G1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 rnm.

Etymology. - The species is named after its type locality (pronounced as "Nan-Aou") andthe name is used as a noun in apposition.

Geothelphusa ilan, new species(Fig. 27)
MateriLll examined. - Holotype - Male, 27.8 by 22.2 rom (NTOU FI0196), ILAN HSIEN: Lotung,colI. U. Hwang, 27.xi.1983.
Paratypes - ILAN HSIEN: Lotung - 5 males (NTOU FI0077), colI. U. Hwang, 27.xi.1983. Tatung - 3 males, 4 females (NTOU FI0078), colI. U. Hwang, 27.xi.1983.
Others - ILAN HSIEN: Sagshing - 5 males, 2 females (NTOU Fl0079), colI. 7.viij,1982; 1 female(NTOU FlO080), colI. Wang, 9.viii.1989. - Tatung Jiuntow - 1 male (NTOU FI0207), colI. J.Y. Shy& W.L. Tsay, 3.ix.199~; 1 male, 2 females (NTOU FI0208), colI. J.Y. Shy & W.L. Tsay, 7.ix.1993.
- Tatung Shnagtsukeng - 9 males (NTOU FI0188), colI. lY. Shy & C.S. Heh, 30.i.1994.

Diagnosis. - Carapace smooth with fine concave pits. Cervical groove distinct; frontal
margin divided into 2 lobes. Anterolateral region with fine striae. Anterolateral crista distinct,
lined with low, small granules, without epibranchial tooth. Distance between tip of maleabdomen and anterior margin of stemite 4 long, about 2.0 times length of stemites 1-3. Fingers
of chela forming oval gape when closed. Lateral margins of male abdominal segment 6
slightly convex. G1 moderately curved outwards, outer proximal margin with a tooth, innerproximal margin dilated; terminal segment moderately curved outwards, spinules lining distal
one-fifth (Fig. 27c-e).

Size. - Largest male 28.5 by 22.3 mm (NTOU FlO077);.largest female 30.3 by 23.7 mm(NTOU F10078); smallest mature female 21.9 by 16.4 mm (NTOU FlO079).

Remarks. - This species is allied to G. nanao, but it distinguished by the following
aspects: 1. the carapace is more convex transversely and longitudinally, the carapace widthand length being about 2.0 times and 1.6 times of the carapace length respectively; 2. thecervical groove is distinct and the anterolateral crista is lined with small granules; and 3. thetotal length of the G1 is about 5.8 times that of the termInal segment.
Etymology. - The species is named after the type locality, and the name is used as a noun
in apposition.

Geothelphusa olea, new species(Fig. 28)
Geothelphusa chiui - Hwang & Mizue, 1985: 13 (part), text fig. 8, pI. lIB (nee Geothelphusa
Minei, 1974)

chiui

MateriLll examined. - Holotype - Male, 21.9 by 16.5 mm (NTOU FI0026), TAINAN HSIEN: Nanhsi,
colI. J.Y. Shy & W.L. Tsay, 7.viij,1992.

Fig. 28. Geothelphusa olea, new species. Holotype male, 30.1 by 23.8 mm. a: dorsal view; b: frontal
view; c: ventral view of right G 1; d: dorsal view of right G 1; e: terminal segment of G 1; f: ventral viewof right G2. Scale = 1.0 mm.

Paratypes - TAINAN HSIEN: Nanhsi - 2 males, 1 female (NTOU FI0026), colI. J.Y. Shy & W.L.
Tsay,7.viii.1992.
HSINCHU HSIEN: Peipu, Nanpu - 10 males, 3 females (NTOU Fl0046), colI. J.Y. Shy & K. Lee,
15.vi.I993.
Others - TAIPEI HSIEN: Chungho -1 male, 1 female [ovigerous] (NTOU Fl0041), colI. C.S. Heh,25.vi.I993; 2 males (NTOU Fl0032), colI. J.Y. Shy & C.S. Heh, 26.vi.I993. - Tanshui - 2 males,
1 female (NTOU FI01l8), colI. J.Y. Shy & K. Lee, 30.iv.1993. - Sanchih - 1 male, 2 females (NTOU
FI01l9), colI. J.Y. Shy, 27.vi.1993; 1 male (NTOU FI0120), colI. C.L. Lee, iii.1993.
TAOYUAN HSIEN: Fuhsing, Kaoyih - 1 male, 1 female (NTOU FlO031), colI. J. Y. Shy & K.
Lee, 14.vi.1993.
HSINCHU HSIEN: Hemgshan - 1 male (NTOU Fl0051), colI. J.Y. Shy, 16.vi.1993.
MIAOLI HSIEN: Shanwan, Tahodi - 4 males (NTOU Fl00430, colI. J.Y. Shy & K. Lee, 15.vi.1993.
TAiCHUNG HSIEN: Taipmg, Tsaohu -1 male, 1 female (TMCD 2844), colI. H.C. Liu, 1O.ix.1988;
1 male (TMCD 2847), ~ll. C.H. Wang, 15.x.1992. - Hoping, Tianlemg - 4 males, 2 females (NTOU
FI0039), colI. J.Y. Shy & W.L. Tsay, 19.x.1992; 1 male, 2 females (NTOU FlO040), colI. J.Y. Shy& K. Lee, 29.vi.1993.
NANTOW HSIEN: Tsaotun, Jeoujeoufong - 2 males, 2 females (NTOU FI0028), colI. J.Y. Shy& K. Lee, 25.xi.1992. - Puli - 1 male (NTOU Fl0037), colI. J.Y. Shy & K. Lee, 27.xi.1992. - Shuili- 1 female [ovigerous] (NTOU FlO047), colI. J.Y. Shy & K. Lee, 30.vi.1993.
YUNLIN HSIEN: Kukeng - 1 male, 1 female (NTOU FlO050), colI. P.H. Ho, 5.ii.1992. - Linnei,
Linpei - 1 male, 3 females [1 ovigerous] (NTOU FlO042), colI. J.Y. Shy & K. Lee, 30.vi.1993.
CHIA YI HSIEN: Meishan - 1 female (NTOU FlO049), colI. H. P. Yu, 9.xi.1982; 1 male, 1 female(NTOU FI0033), colI. J.Y. Shy, l.vii.1993. - Chuchi - 2 females (NTOU FlO030), colI. J.Y. Shy &P.H. Ho, 16.xii.1990; 1 female (NTOU FI0038), colI. J.Y. Shy & W.L. Tsay, 15.vii.1992; 1 male, 4females (NTOU FI0034), colI. J.Y. Shy, 21.xii.1992. - Chuchi, Paichii - 2 males (NTOU FlO036),
J.Y. Shy, 19.xii.1992. - Meishan, Yeongshing 2 males, 1 female (NTOU FI0029, colI. J.Y. Shy &H.G. Lai. - Tapu, Tamaopu 3 males, 2 females (NTOU FI0027), colI J.Y. Shy & K. Lee, 28.xi.1992.
CHIA YI CITY: Jiuntou - 1 male, 1 female (NTOU FlO048), colI. J.Y. Shy & H.G. Lai, 3.iv.1992.
TAiNAN HSIEN: Yuching - 19 males, 2 females (NTOU FI0044), colI. H.P. Yu, 12.xi.1983.Nanshi - 1 female (NTOU FlO045), colI. J.Y. Shy, 7.viii.1992.
KAOHSIUNG HSIEN: Sanmin, Minchu - 1 male (NTOU Fl0035), colI. J.Y. Shy & W.L. Tsay,
6.viii.1992.

Diagnosis. - Gastric region of carapace smooth. Anterolateral region slightly rough,branchial region flat. Epibranchial tooth faint, anterolateral crista distinct, lined with small,
low granules. Fingers of chela forming wide, oval gape when closed. G 1 slightly curved
outwards, base of subterminal segment narrow, total length about 4.1 times width of base,outer proximal margin with a tooth; terminal segment moderately curved inwards, total length
about 7.5 times of terminal segment length (Fig. 28c-e).
Coloration. - Carapace and ambulatory legs greenish-gray to yellowish-green. In younger
individuals, the ambulatory legs are covered with scattered darker coloured stripes.

Size. - Largest male 35.4 by 28.2 mm (NTOU FlO031); largest female 36.2 by 27.9 mm(NTOU F10030); smallest mature female 21.2 by 17.0 mm (NTOU FlO044).

Remarks. - This species is allied to G. albogilva, but it distineuished by the following
aspects: 1. the anterolateral crista is distinct, lined with small granules and has a small
epibranchial tooth; 2. the base of the G 1 subterminal segment is narrow and the length of

the synovial membrane is about 4.4 times that of the width; and 3. the distance between the
tip of the male abdomen and anterior margin of sternite 4 is about 1.4 times of sternites 13 (about 1.7 times in G. albogilva).
Etymology. - The name is derived from the Latin for olive, alluding to the colour of thecarapace. The name is used as a noun in apposition.

We are greatly indebted to thank Mr. C. H. Wang of the Taiwan provincial Museum formany valuable suggestions- and providing us with material. Mr. P. H. Ho, Miss K. Lee andMiss W. L. Tsay oil'National Taiwan Ocean University, Department of Fishery Science,
Crustacea Laboratory for helping us in field collections and taking some of slides. Mr. H.
C. Liu of National Tsing Hua University, Graduate School of Life Sciences, providing uswith the material. Dr. Keiji Baba kindly checked on the status of the types of G. chiui, andhe and Dr. Masatsune Takeda provided information on Minei's types. This present report
is a contribution from a research grant on the Decapoda Crustacea of Taiwan supported by
the National Science Council, Republic of China.

Alcock, A., 1910. Brachyura I. Fasc. II. The Indian Freshwater Crabs - Potamonidae. Catalogue ofthe Indian Decapod Crustacea in the collection of the Indian Museum. Calcutta, pp. 1-135, PIs. 1-14.
Balss, H., 1937. Potamoniden (Dekapoda Brachyura) der Philippinen und des Malayischen Archipels.
Intern. Rev. d. ges. Hydrobiol. a. Hydrogr., 34(3-5): 143-187.
Bott, R., 1967. Potamiden aus Ost-Asien (Parapotamon
de Man, Sinopotamon
n. gen.,Candidiopotamon n. gen., Geothelphusa Stimpson) (Crustacea, Decapoda). Senckenbergiana bioi.,
Frankfurt, 48(3): 203-220.
Bott, R., 1968. Parathelphusiden aus Hinterindien (Crustacea, Decapoda, Parathelphusidae).
Senckenbergiana bioi., Frankfurt, 49(5): 403-422.
Bott, R., 1969. Fliisskrabbenaus Asien und ihre Klassifikation.(Crustacea,Decapoda).Senckenbergianabioi., Frankfurt, 50(5/6): 359-366.
Bott, R., 1970. Die Siisswasserkrabben von Europa, Asien, Australien und ihre Stammesgeschichte.
Eine Revision der Potamoidea und Parathelphusoidea (Crustacea, Decapoda). Abhand. Senckenberg.
Naturf Gesell., Frankfurt, 526: 1-338, PIs. 1-58.
Man, de, 1. G., 1914. Note sur quelques crustaces decapodes brachyures terrestres et d'eau douceappartenant au Musee Civique de Genes. Ann. Mus. Civ. Stor. Nat., 6: 128-135.
Haan, W., De, 1833-1835. Crustacea. In: Fauna Japonica auctore Ph. Fr. de Siebold, 1833: 1-24; 1835:
23-64.

Hwang, J. J. & K. Mizue, 1985. Fresh-water crabs of Taiwan. Bull. Fac. Fish., Nagasaki Univ., 57:
1-21, PIs. 1,2.
Kemp, S., 1913. Zoological Results of the Abor Expedition, 1911-1912. XX. Crustacea Decapoda.
Rec. Ind. Mus., 8: 289-310, PIs. 17-21.

~-...

.].~.
__ .._ ..

..-..... /._/,'----~:"~
'...

.

,

I'
/

'"

_. -...~-. -- --(.

\

,,

,

,

,,

o

••

H L

''i!

Fig. 29. Map of Taiwan showing main districts.: (TPl) Taipei City; (TP2) Taipei Hsien; (KL) Keelung
City; (TY) Taoyung Hsien; (HC) Hsinchu Hsien; (ML) Miaoli Hsien; (TC) Taichung Hsien; (CH)
Chunghwa Hsien; (NT) Nantow Hsien; (YL) Yunlin Hsien; (CY) Chiayi Hsien; (TN) Tainan Hsien;(KH) Kaohsiung Hsien; (PT) Pingtung Hsien; (IL) l1an Hsien; (HL) Hwalien Hsien; (TT) Taitung
Hsien; (LT) Lutao; (LY) Lanyu.

0<
"

..'r'
... ;....

\ ...J. .
'/.----

.

~:~

.-...

:'

:'

~.
-- _ ... ,,'

'.

.

,

.•.-••.•...... \...

.-J..,···'tr.~-"..::-;'.

o

••

~...
'

[',

''il

.-/*
[',

i,

,g
"

"

o

0

l0000
0 I

v~~

...__ ..

?;':;:~)
/l
.•.•

.....
....
··..t..·.
.•... __ . .
",....·--.--1,."------\.•.
.-.
.•........•..-/••.'.---,-/.:.:\.•....
'... '..

---

,<;t

Q

.'

Fig. 31. The distributions of seven species of freshwater crabs in Taiwan.

o : G. dolichopodes;

6:

G. eucrinodonta;© : G. eurysoma;:"\··.,0·· .•

o

o

..

:

,

i 0\7 •.._••••.

.

:\.~ ••• y ••:.

\ ..'J

.'

o0 )
o qi
~...

~:::

o

.:,

.,..'

.

0

.... "

:'

:'
..••

'"
'.-

•• _- --"__ ••• -r".
,.... -----)".-.

0'·-·'
D D D.......

....

'"

0*

..".-.--.-.;.

D....:.

0.(;········,-/

o

.

,"

,.S···""

./!

'\._.
"-', ".

Do',:'.'

Fig, 32, The distributions of seven species of freshwater crabs in Taiwan,o : G, lutao;o : G, nanao;

6 :G, miyazakii;© : G. monticola;

0 : G. olea;

V : G. takuan.

-(:(: G. nanhsi;

.

:J~-~/-- ... - .. -- -- --".

"0

o

''il

Fig. 33. The distributions of seven species of freshwater crabs in Taiwan.

o : G. tali;

6 : G. taroko;© : G. tawu;

Minei, H., 1973. Potamoid crabs of the Ryukyu islands, with description of five new species (Crustacea,
Decapoda, Potamoidea). J. Fac. Agric., Kyushu Univ., 17: 203-226.
Minei, H., 1974. Potamoid crabs of Taiwan, with description of one new species (Crustacea, Decapoda).
J. Fac. Agric., Kyushu Univ., 18: 239-251.
Miers, E. J., 1886. Report on the Brachyura collected by H.M.S. Challenger during the years 18731876. Rep. Voy. Challenger, 17(2): 1-362, PIs. 1-29.
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"

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and Parathelphusidae(Crustacea:

Decapoda:

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Rathbun, M. J., 1898. Description of three new species of freshwater
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crabs of the genus Potamon.

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Shen, C. J., 1932. The Brachyuran Crustacea of North China. Zoologica Sinica, (A), Invertebrates ofChina, 9(1): i-x, 1-320, PIs. 1-10, 1 Map.
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